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SMITHSONIAN
MISCELLANEOUS COLLECTIONS

VOL. 7

TOLL)

“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—JAMES SMITHSON

(Pustication 4134)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1953

The Lord Baftimore Press

BALTIMORE, MD., U. S. Ae

The Smithsonian Miscellaneous Collections series contains, since the
suspension in 1916 of the Smithsonian Contributions to Knowledge,
all the publications issued directly by the Institution except the An-
nual Report and occasional publications of a special nature. As the
name of the series implies, its scope is not limited, and the volumes
thus far issued relate to nearly every branch of science. Papers in
the fields of biology, geology, anthropology, and astrophysics have
predominated.

LEONARD CARMICHAEL,
Secretary, Smithsonian Institution.

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CONTENTS

BARBER, HERBERT SPENCER. North American fireflies of the
genus Photuris. With preface and notes by Frank A. McDer-
mott. 58 pp., 3 figs. Nov. 27, 1951. (Publ. 4051.)

WETMORE, ALEXANDER. Additional forms of birds from Colom-
bia and Panama. 11 pp. Sept. 25, 1951. (Publ. 4052.)

SHAW, F. R., and Suaw, M. M. Relationships of certain genera
of fungus gnats of the family Mycetophilidae. 23 pp., 45 figs.
Dec. 27, 1951. (Publ. 4053.)

WETMORE, ALEXANDER. A revised classification for the birds of
the world. 22 pp. Nov. 1, 1951. (Publ. 4057.)

EISENMANN, EUGENE. Annotated list of birds of Barro Colorado
Island, Panama Canal Zone. 62 pp. Feb. 7, 1952. (Publ.
4058.)

Emerson, WiLL1AM K. The scaphopod mollusks collected by the
First Johnson-Smithsonian Deep-Sea Expedition. 14 pp., I pl.
Feb. 26, 1952. (Publ. 4059.)

CrarkE, J. F. Gates. Host relationships of moths of the genera
Depressaria and Agonopterix, with descriptions of new species.
20 pp, GO pls. 2pr.i23, 1952. \( Publ. 4083.)

Snoperass, R. E. The sand crab Emerita talpoida (Say) and
some of its relatives. 34 pp., 11 figs. Apr. 15, 1952. (Publ.
4086. )

Azpot, C. G. Precipitation and temperature in Washington,
D. C., for 1951 and 1952. 5 pp., 2 figs. Mar. 18, 1952. (Publ.
4087.)

Assot, C. G. Periodicities in the solar-constant measures. 31 pp.,
6 figs. May 28, 1952. (Publ. 4088.)

Assgot, C. G. Important interferences with normals in weather
records, associated with sunspot frequency. 3 pp., I fig. May
20, 1952. (Publ. 4090.)

Drucker, Puitip. Two aboriginal works of art from the Vera-
cruz coast. 7 pp., 3 pls., 1 fig. Aug. 26, 1952. (Publ. 4091.)
KnicuHT, J. Brookes. Primitive fossil gastropods and their bear-
ing on gastropod classification. 56 pp., 2 pls., 10 figs. Oct. 29,

1952. (Publ. 4092.)
(v)

15.

16.

17.

18.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

Coorrer, G. Arruur. New and unusual species of brachiopods
from the Arbuckle group in Oklahoma. 35 pp., 4 pls. Sept. 23,
1952. (Publ. 4093.)

Loesticu, ALFRED R., Jr., and TAppAN, HELEN. The forami-
niferal genus Triplasia Reuss, 1854. 61 pp., 8 pls., 11 figs.
Sept. 9, 1952. (Publ. 4094.)

Assot, C. G. Solar variation and precipitation at Peoria, Illi-
nois. 18 pp., 8 figs. Sept. 3, 1952. (Publ. 4095.)

Mirtteman, M. B. A generic synopsis of the lizards of the sub-
family Lygosominae. 35 pp. Nov. 4, 1952. (Publ. 4096.)

Gazin, C. Lewis. The lower Eocene Knight formation of west-
ern Wyoming and its mammalian faunas. 82 pp., 11 pls., 6 figs.
Dec. 9, 1952. (Publ. 4097.)

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NORTH AMERICAN FIREFLIES OF THE
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HERBERT SPENCER BARBER .
WITH PREF ACE AND NOTES BY FRANK A. McDERMOTT

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Beat “CITY OF WASHINGTON
"PUBLISHED, BY THE SMITHSONIAN INSTITUTION
\ ; NOVEMBER 27, 1951

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 1

Thomas Lincoln Casey Fund

NORTH AMERICAN FIREFLIES OF THE
GENUS PHOTURIS

BY
HERBERT SPENCER BARBER
WITH PREFACE AND NOTES BY FRANK A. McDERMOTT

(PUBLICATION 4051)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 27, 1951

The Lord Baltimore Press :
BALTIMORE, MD., U. & As oii

i
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PREFACE

At the time of his death, on June 1, 1950, Herbert Barber had
nearly completed the manuscript of a monograph on the North Ameri-
can fireflies of the genus Photuris. Subsequently, I was requested by
Dr. E. A. Chapin, curator of the division of insects, U. S. National
Museum, to review the manuscript with a view toward putting it in
shape for publication. I have done this with mixed feelings of wonder
at the amount of field work involved (not always in the easiest places
for such studies), of admiration for Barber’s persistent checking and
rechecking of observations, and the logical deductions he has drawn,
and of regret that he was not able to complete the work to his own
satisfaction. Actually, the monograph as he wrote it is so nearly
complete that little more than the correction of a few obvious typo-
graphical errors and the change of an occasional word or punctuation
mark has been made in the text. His pencil sketch diagramming the
flashes of the males of the various species of Photuris has been re-
drawn for reproduction, with the addition of those species he describes
but did not include in his sketch.

The beetles of the family Lampyridae are almost unique among
insects because of the ability of most species to produce light, a func-
tion limited to only a few other insects, although widely distributed
among marine forms. As in most other insects, the family has been
divided into a large number of genera, one of the most distinctive
of which is Photuris, limited at present to New World species, and
being more or less replaced in the Old World by the genus Lucziola,
of somewhat similar characteristics.

The generic name Photuris was first used by Dejean (1833), estab-
lished by LeConte (1852), and subsequently used by Lacordaire
(1857), Olivier (1886), and others, for species presumably falling
naturally into this classification. The vagaries introduced by several
authors have been sifted by Mr. Barber, and the details are given
in the text of this monograph. Barber has done a beautiful piece of
work in unraveling the tangled skein of nearly a score of morpho-
logically very similar species, many with adjacent but overlapping
habitats, and with distinct mating habits. He says, “All structures,
even those of the male genitalia, appear identical in our numerous
species.” His manuscript refers to sketches of the aedeagus of
Photuris frontalis, which he uses as typical, but his sketches have not

iii

iv PREFACE

been found, and I have substituted for them sketches of this struc-
ture from Photuris lucicrescens from Delaware. In Barber’s segre-
gated set of 19 species and varieties, 10 show the aedeagus extruded,
and except for size there is no observable difference in the different
species; the aedeagus of Photuris jamaicensis, sketches of which
have been kindly lent me by Dr. John B. Buck, is also apparently
identical with that of Barber’s species.

That Barber was able to recognize his species in dried specimens,
when he had not seen the flashing conduct, was demonstrated to me
when I submitted to him a series of five vials containing specimens
collected around Wilmington, Del., each vial representing a different
flash ; for four of the vials he told me correctly the type of flash after
a few minutes examination with a lens; the fifth contained specimens
having a flash with which he was not familiar, and which were
probably abnormals.

This work of Mr. Barber may stem ultimately—aside from his
general interest in the Lampyridae—from a conversation between
him, Dr. E. A. Schwarz, and myself in 1910, when I was studying
the relation between light emission and mating habits of the fireflies.
Photuris pensylvanica was mentioned, and Dr. Schwarz remarked
on the enormous numbers in which it occurred in Panama. I asked
if it were the same species, and he replied something to the effect
that it was the same species from Massachusetts to Panama, and then
added that “some day somebody is going to split that thing up.” This
Mr. Barber has done with infinite care and persistent checking. That
portions of the picture are still confusing cannot be denied, but it
is very evident that what was long considered to be a single species
is undoubtedly a complex of many morphologically closely similar
species with quite distinct habits, habitats, and mating behavior. If
these have to be considered “physiological species,” so be it.

That this condition is not peculiar to Photuris is indicated by an-
other case among the Lampyridae described to me by Mr. Barber in
June 1947. While collecting specimens of the supposed Lecontea
(Pyractomena) lucifera (Melsheimer), near Washington, he en-
countered a species giving a single bluish flash instead of the twin-
kling 5-component flash previously ascribed to lucifera, but upon in-
spection of his vials the next day he concluded he had mixed his speci-
mens, as all those he had taken were apparently identical. On his next
trip he very carefully segregated the 1-flashers from the 5-flashers,
but upon inspection he was again unable to tell one from the other
by general appearance. In this case the aedeagi proved to be different,

PREFACE = ON,

but the question still remains as to which of the two is the one origi-
nally called lucifera by Melsheimer.

While the original manuscript of this monograph was apparently
prepared by Mr. Barber about 1929, his interest in the problem con-
tinued unabated, and at least two species were defined after that time.
His letters to me from 1926 to 1929 indicate the development of the
ideas given in the monograph, and later correspondence gave evidence
of the confirmation and extension of his observations, and also re-
corded his difficulties with cabinet specimens. Mr. Barber had started,
about 5 years ago, an extensive review of the taxonomy of the
Lampyridae, which work was most regrettably interrupted by his
death. His notes show a complete grasp of the difficulties involved,
an accurate and broad knowledge of the literature, and the modern
concept of a species as a dynamic unit, a breeding population. Per-
haps some quotations from his notes made in the course of his work
on this revision may not be amiss:

Dated February 14, 1945:

The writer’s belief that each species is an isolated self-perpetuating popula-
tion, limitless in individuals by past and future generations, and that our tax-
onomy must correctly interpret these natural species which contrast so hope-
lessly with the customary “taxonomic” species, has combined with his inability
to apply the available names to his samples of “natural” species, to discourage
completion of manuscripts.

Undated, probably 1944:

If, however, a collector seeks the luminous species when they are active,
distinguishes the signals of the several species which may be in their nuptial
flight, and the peculiar flashes emitted by the opposite sexes of each, and col-
lects individuals which emit a particular type of flash, his samples thus assembled
and segregated will more correctly represent the unit species he has observed.
If, also, the observer selects convenient undisturbed localities, such as upland
fields, woods, river banks, marshes, etc., in which he can repeat his observations
in successive seasons and years, he may find that each peculiar habitat has its
sequence of species peculiar to it, their larvae present most of the year, the
adults active for only a few days at the correct season, except the unpredictable
abnormal individuals who leave no progeny. Repeated verification of observa-
tions is essential.

As late as September 1949 he wrote to me—

This problem (the species of Photuris) is far more complicated than you
think, and we are still far from the truth. Taxonomy from old mummies which
fill collections is a misguided concept. It leads to the misidentification of rotten
old samples in collections. How these poor fireflies would resent being placed
in such diverse company—among specimens of enemy species—if they were
alive and intelligent! What contempt they would feel for the “damned
taxonomist.”

V1 PREFACE

Barber’s own field observations covered, for the most part, a rather
limited geographical range, roughly within a radius of about 100 miles
from Washington, D. C., and to judge from the species from Wis-
consin, Cape Breton, and elsewhere it is quite possible that a num-
ber of other types of flashing conduct may be found in areas outside
of those he covered. Only future work can show how general the
distribution of his species may be and how much overlap in range
may occur.

The practical mind may ask, “Of what use is such a study?” To
which we may reply with Faraday’s famous retort to Gladstone: “Of
what use is a newborn babe?” Aside from the basic “increase and
diffusion of knowledge among men,” we can never tell when, where,
and how a given observation may be of practical importance. The
chance observation of a bacteriologist a quarter of a century ago,
that bacteria did not grow in the presence of a mold, is the basis of
the multimillion-dollar antibiotics industry of today. The possible
importance of the Lampyridae as predators against agricultural pests
has barely been touched upon (see p. 2 of this monograph, and the
writer’s “Common Fireflies of Delaware,’ Wilmington, 1948). In
any event, such a study as this of Mr. Barber’s on Photuris is, as
Emerson says of beauty, “its own excuse for being.”

Frank A. McDermott.
Wilmington, Del.
December 4, 1050.

Thomas Lincoln Casey Fund
NORTH AMERICAN FIREFLIES OF THE
GENUS PHOTURIS
By HERBERT SPENCER BARBER}
Division of Insect Identification

Bureau of Entomology and Plant Quarantine
United States Department of Agriculture

CONTENTS

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Notes on some general characters of North American Photuris....... 37
Results of the examination of Barber’s specimens.............++++e00- 43
PTETALTIRE FOILCO ercncss yeteoeare ere eide core e oiern Mahe Canvey eio a ela jase: ays) eyecs nara ayer esierals 56

INTRODUCTION

Although it has been argued that so-called “physiological species”
should not be given distinctive names, inasmuch as they cannot be
identified from average cabinet specimens, this attitude is opposed to
the objectives of the study of natural history. The demand for visible
external characters by which species may be “identified” and the ex-
altation of this principle as a standard of specific value have already,
within the memory of most of us, broken down before the newer
standard (useful in many groups but not universally so) based upon
internal or reproductive organs. Species being biological units com-
posed of populations reproducing their kind and supposedly isolated
from other species by barriers of some kind, it behooves the student
to find the characters by which they may be recognized. If these char-
acters are external and “structural” in the old sense, the investigator

1¥For a biographical sketch of Mr. Barber, see Proc. Ent. Soc. Washington,
vol. 52, pp. 259-269, 1950.—EpiTor.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 1

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

is lucky. But if all the customarily used structural characters of
shapes, sizes, and colors are variable within all of several allied
species which are distinct in ecological habitat, time of maturity,
habits, and courtship behavior, the student must find new standards
or abandon his study. In the face of general opinion among sys-
tematists, which opinion has itself become standardized, the tempta-
tion is to take the latter course. But it is a pity that so few systema-
tists realize that the only fundamental object of naming species is, in
the ideal, to produce a system by which records of observed facts
about species may be indexed so that the students of insect economy,
behavior, anatomy, genetics, etc., as well as the systematist, may
assemble and sort the desired data. Surely the confusion will be inex-
tricable if only those species that chance to display some “structural”
character receive distinctive names.”

In some groups specialization in structure seems to have occurred
without apparent specialization in habits; in other groups structures
remain practically identical but habits have become distinct; and in
still others the exoskeletal variation within members of a brood may
render ordinarily used characters useless, or plasticity of habits may,
by accident, accompany temporary development of a differential
habitus. Hybridization may be so common as to unite similar species
into a variable or even homogeneous population, or may be so rare
that fertile offspring, fit for reabsorption into either of the self-
perpetuating species, are as infrequent as in mules.

In the genus Photuris individual variation is so common that the
following studies have yielded few characters besides certain gen-
eralized differences of color, size, and, in a few cases, proportion ;
but since these are connected with habitat and habit distinctions, they
must serve until better diagnostic characters are found. All struc-
tures, even those of the male genitalia, appear identical in our
numerous species.

Although Photuris larvae are general predators on snails and soft-
bodied insects and may be of some economic value as enemies of cut-
worms (Hess, 1920),° the specialization of different but hitherto con-
fused species to different and particular types of breeding ground
indicates diversity in the preferred prey. Records of observations on

2 Dr. Ferris has published similar views (Ferris, G. F., The principles of sys-
tematic entomology, p. 48, Stanford, Calif., 1928). For a further discussion see
Mayr, Ernst, Systematics and the origin of species, New York, 1942 (1949),
particularly p. 20, “What is a taxonomic character ?”—McD.

’ Names and years in parentheses refer to the Literature Cited at the end of
the monograph.—McD.

NO, I FIREFLIES OF THE GENUS PHOTURIS—BARBER 3

feeding habits are fragmentary and cannot now be associated with
particular species, but it should be obvious that marsh-inhabiting
species could have little influence on a cutworm infestation in an
adjoining field, whereas an abundant upland form, such as pyralomi-
mus, described below, might be an important enemy.

OBSERVATIONS ON ADULT BEHAVIOR

Since the writer’s interpretation of our Photuris fauna as he has
observed it differs from that of students of fireflies who rely upon
dried specimens and upon the standards of the older taxonomists, he
begs for a moment that readers imagine themselves sharing with him
a few selected experiences that have forced great changes in his belief
as to what constitutes a species.

1. A hilltop field of grass bordered by woods in Rock Creek Park,
D. C., early in June—Photuris have just appeared here in the past
few days, and on this evening they are flying in numbers over the
field but not in the woods. Many are flashing in the gathering dusk
as they fly a few feet above the grass, and only two types of flash
are apparent in the air, the commonest being a series of about six
very quick flashes in less than a half second, of not great brilliance.
These are all males flashing their signals, hoping for answering flashes
from prospective mates. Rarely one may see such a response in the
short grass—a brief, less brilliant, single glow of about a third of a
second duration—and observe the quickened repetition of the male’s
signals as he approaches in a long oblique descent. This female flash
appears seldom in the air. Green leaves and the fingers are held over
the bulb of a small flashlamp by the observer, concealed among
foliage, and an attempt to mimic the female flash is made immediately
following the flash of a nearby male. He comes rapidly to the hand
and is caught ; other males have seen the mimic of the female’s flash
and are coming also, so that the collector may catch half a dozen with
the hand without moving from the edge of a concealing bush. A
steady light does not attract, but frightens the males away. Another
type of flash is occasionally seen as we ramble about, but it is the
short, frequent, but very irregular flash of disturbed individuals,
usually females, whose agitation is visible in abnormal functioning
of the light organ.

This species the writer identifies doubtfully as versicolor Fabricius,
which was described in 1798, without more definite locality than
North America, from a specimen received from Mr. Hirschell and
has since been incorrectly suppressed as a synonym of an earlier
given specific name.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

2. A rocky, heavily wooded island in the gorge of the Potomac
late in June.—At the downstream end an alluvial deposit subject to
freshets supports a dense growth of maples bound together by a canopy
of wild grapevines, hiding the stars; underneath, a few sandy freshet
channels can serve as paths. The great wood interior is filled with
innumerable flashes of greenish-white light, and at first there seems
no variation in the flashes. Each firefly appears to give a single short,
very bright flash for each second that it flies, and all those flashing
are males. An occasional slightly different flash on foliage or ground
is investigated and discloses the presence of females, which have pre-
sumably mated and are not at all interested in the self-assertive males.

Two or three times during the preceding winter and spring the
floods have swept for days, roaring between the trees and among the
sandbars, bringing logs and smaller driftwood, which lie in masses
where the trees chance to hold them. Other species of fireflies appear
discouraged by such abuse of their breeding ground, but before the
firefly season comes, the glowworms of this form are abundant in and
about these masses of river drift, above and in the immediate vicinity
of which the males later fly in numbers. Occasional individual adults
are to be seen in every few hundred feet of river forest in June, but
these are supposedly strays maturing where they were left as larvae
by the water. The spring freshet of 1928 washed out the glade in
which the species was watched the two preceding years, but it left an
accumulation of drift on some logs 50 feet to one side. Few of this
species were seen where formerly abundant, but they later became
numerous about the driftwood. This species is herein named potomaca,
p. 28.

3. Crossing the current to the Virginia shore, we see the same
species in fewer numbers in the fringe of trees on the bank, but in
the field behind are a few belated males of the flicker-flash species
above described (versicolor Fabricius?). The path crosses the small
neglected field and dips into a damp hollow carrying the drainage
from Black Pond and bordered with scattered willows, beyond which
the ground rises a few feet to a terrace upon which low alders grow.
Then there is another narrow grassy strip and the wooded rocky hill-
side rises abruptly. The willows and low vegetation along the slug-
gish stream are glittering with myriads of flashes, of almost the same
short duration and interval as the greenish lights we have just left
in the river forest, but these are faintly orange instead of greenish,
and slightly slower, about three flashes in 4 seconds. The samples
caught are all males, but are smaller in size, differently colored, and

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 5

have slightly more elongate antennae. This species is herein named
hebes, p. 34.

4. A few steps farther, the path enters the alders and immediately
a very different type of flash confronts us. Poising almost motion-
less in the air, its light begins dim, grows steadily to great brilliance
and dies abruptly, to reappear a quarter or half minute later as the
firefly poises a few feet distant and again remains illuminated for
from I to 2} seconds. All these are, as before, males, but they are
larger, broader, and much paler in color. Their females are found
demurely about their business of seeking food, for the female Photuris
eats other fireflies ; but since no courtship is observable they are sup-
posed to be already mated individuals no longer interested in the
surrounding lights. This species is herein named lucicrescens, p. 33.

5. Drive 15 miles to the tide marsh of the Anacostia River, and
even though the hour is midnight Photuris of several species are still
flashing. The long crescendo flash just described is conspicuous in the
bushes bordering the marsh, and in the treetops is a very short, bright
flash, almost an explosion of light, at 4- or 5-second intervals. Sam-
ples of this species we cannot reach in its normal flight. But over the
level tops of the tall, rank grass of the marsh another very different
flash greets us—an instantaneous explosion of light followed immedi-
ately after an extremely short, dark interruption by a protracted
brilliant light lasting I to 2 seconds, with the end perceptibly dimin-
ished in intensity. We wade into the deep grass and ooze and catch
samples. They are not half so large as the crescendo-flash species on
shore, and some have wing covers pale except basal remnants of the
brown vittae. Certainly it is the only species seen tonight to which
the original habit notes and description of pensylvanica (original
spelling of specific name), published by De Geer more than a century
and a half ago, can be applied. While emitting this double flash the
male (for no females are visible to us) poises in his flight over the
grass tops, dips slightly and rises, describing little U-shaped curves
of light, the finish a little higher than the first flash. He must watch
for his bride’s answer straight beneath, since marsh grass stands ver-
tical at this season and cannot be seen through obliquely. But his be-
havior is the result of instinct instead of reason and reflects an
immensely old specific adaptation to this particular ecologic environ-
ment. No females can be found while we walk forward, but if we turn
and force our way backward through the grass their annoyed flashes
deep in the disturbed grass or on the surface of the ooze permit their
capture in numbers. In the vial used to preserve these females I find

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

a minute fish (Umbra). Wasa female eating a fish when caught? No
other debris is in the vial.

6. Dense Baccharis bushes on a sand spit joining a wave-eroded
bluff surmounted by oaks and pines, overlooking the brackish water
of an arm of the Chesapeake early in July—A warm evening breeze
sways the bushes and low among them, or rising in their lee, fly
moderate numbers of a small firefly emitting short, abrupt, faintly
orange flashes at intervals of about 3 seconds. Specimens caught re-
semble the small, willow-swamp form (hebes, above), and the long
double-flash species of the fresh-water tide marsh (pensylvanica),
but they seem to have larger eyes and shorter antennae than these
others.

Again, a small salt meadow near the mouth of the Potomac estuary,
in front of pines, hollies, oaks, Myrica, Baccharis, and Iva bushes,
in successively more frequently inundated tidal shore line than the
wetter salt marsh—Among these bushes and straying among the
nearby grass tops appear short, slightly orange flashes at 2- to 3-
second intervals, but the insects keep well down where the shore
breezes do not blow them away from their native habitat, thus con-
trasting strongly with the other species visible in the woods.

The small size of the firefly and its feeble flash resemble those of
hebes, but the preserved samples differ in that this salt-marsh species
shows larger eyes, shorter and stouter antennae, a black labrum, and
a broad, black, midpronotal vitta. The ancestors of this species hav-
ing for ages past held their place among the shore bushes against
breezes, the generation now under observation flies low among the
sheltering bushes undisturbed by a mild wind which scatters and
forces down the flight of hebes. We shall later (p. 35) name this
new form Photuris salinus.

7. Varying from year to year with the earliness or lateness of the
season, the flicker-flash species (versicolor) appears in the above-
described field at Black Pond about the middle of May and has be-
come relatively scarce by the second week of June, when it is replaced
by a slightly smaller form whose males, when not disturbed, appear
to have two distinct types of light signals. This form seems to origi-
nate from the swampy ground among the willow and alder clumps
some two weeks before the larger species, lucicrescens, and the smaller
one, hebes, above discussed, begin to be seen. The behavior of this
intermediate species (if it be but one form with two habits) will be
variously interpreted according to preconceived notions, but requires
record here.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 7.

As the sky colors fade to gray, the first sharp, greenish-white flashes
appear in the clumps of bushes and on their darker eastern sides, con-
trasting strongly with the feebler orange flashes of the few early
males of Photinus scintillans, which almost immediately cease their
activity. As dusk deepens, the Photuris become numerous, the very
short, sharp flashes being emitted at intervals of from 3 seconds, on
a pleasant evening, to perhaps 10 seconds, if it is cool and there is
heavy dew. In the latter case they soon cease to fly and their slower
flashes emanate from males resting on foliage in slightly more sheltered
situations. But from time to time there appear among them males
flying slowly over grass or bushes, or even resting on foliage if it
has become cool, and emitting long, tremulous flashes, less intense
than the commoner sharp flash, consisting of perhaps Io to 20 pulsa-
tions, and lasting about a second. Within a quadrant of perhaps 50
yards’ radius from the same point of observation, these long tremu-
lous flashes may appear, followed by others, becoming more and more
numerous, the shorter flashes disappearing until for a few minutes
the long flashes dominate. This phenomenon suggests either that
another species has temporarily become active, as the writer has often
observed with certain species of Photinus, or that a contagious emo-
tional exuberance has changed the behavior of those males formerly
emitting the short flashes. Samples of the producers of each type of
flash are not distinguishable, as in the case of Photinus above alluded
to, and are hereinafter (p. 31) described as but one species, tremulans.

8. Late in July the swampy forest bordering the Patuxent River at
Priest’s Bridge, Md., is visited. As on previous visits during the pre-
ceding three weeks, only one species of Photuris (lucicrescens) seems
to be active, displaying its long, crescendo flashes, but the numbers
are now much reduced, and the flash appears shorter compared with
our half-second pendulum, used for estimating duration of flash and
of dark interval. The light appears to last from three-fourths second
to about one and one-half seconds. The treetops are watched for
the very short flashes seen elsewhere, but none are seen there now,
nor were they seen on previous visits. We return along the road to
the Capital, stopping when colonies of fireflies are seen. Photinus
pyralis having ceased its activity at an earlier hour, no flashes are
seen except about trees bordering wet spots in the hollows, usually
swampy courses of small streams. Two such places show only the
crescendo flashes, but about 6 miles west of Priest’s Bridge we first
see numbers of the very short explosions of light in the air about
the tree tops. A gust of wind disturbs the fireflies, and one comes
down among the lower branches flashing at about 5-second intervals,

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

very bright and short, about like the one-tenth-second camera shutter
held against a light. When almost within reach the strong spotlight
beam is abruptly thrown on him, and the net brings him to hand. No
chance this time to have netted the wrong firefly, but he looks no
different from those taken in their long crescendo flash. No more
come down, and we must give up and go home.

Why were none of these flashes seen at Priest’s Bridge or at two
other stations? Why do both types of flash occur here and at some
other places?

Various answers will satisfy various persons, but no one knows.
Envy the bats their wings? With them we might follow single speci-
mens through their evening’s activities and see if they change their
flashes.

By the first week in August the firefly population of the wooded
island, the alluvial field, the willow-lined marshy stream, and the
alder bushes near Black Pond has changed. A few belated females
and an occasional male of the large crescendo-flash species (luci-
crescens) are mixed with larger numbers of the short-flashing, smaller
form (hebes) but are no longer confined to the restricted areas as
observed in June. Abnormals appear in all populations, and these
late-issuing individuals may have been lacking in some of the factors
inducing early transformation or fertilization, and the resulting rest-
less dispersal flights may have carried them far beyond the preferred
breeding ground. The whole impression is that of meaningless varia-
tion, and doubts of specific significance are inevitable under such
conditions. In the tidal marshes the little double-flash species (pensyl-
vanica) has vanished, and from the shore forests strays of other
species, most of them females, have wandered out over the marsh
where they mingle with surviving individuals of a small Pyractomena
and several small species of Photinus. Here again one can see only
chaos in their behavior, but next year at the proper time and place
the new generations will court their mates in a similar manner. Spe-
cific flashes will win specific answers, leading to reproduction. Per-
haps we may learn that the manner of flashing is a barrier to possible
intermixing of species. Perhaps the late-season abnormals are mix-
tures. Must we then ignore the differences in the early-season broods?

g. Through the kind interest of friends, observations and well-
preserved samples of Photuris are available from the vicinity of
Winona, Minn., where three apparently distinct species were encoun-

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 9

tered on the evenings of July 6 and 8, 1926, by Miss E. Myers and
B. Boland. Two localities were examined: The first, which was
rather dry land, 10 miles west of Winona on the road to Stockton,
yielded 13 males of a form (versicolor var.?) whose males emitted
five short, greenish flashes as fast as one could count, at perhaps half-
minute intervals, while flying 2 or 3 feet above the tips of the tall
weeds, and four males (caerulucens) that emitted a slow, blue-green
flash of about 1 second’s duration, whose light was dimly visible after
the end of the flash. Flashes of the latter species were seen in much
greater numbers in more open pastureland nearby and over trees.
This latter species was taken in series (30 specimens) at the second
locality in Wisconsin between Dodge and Bluff Siding, 10 miles north-
east of Winona, producing the bluish-green, 1-second flashes over
damp ground near a tamarack swamp. With it in almost equal num-
bers (24 specimens preserved) flew a slightly smaller but otherwise
similar species (aureolucens) that emitted single, short, orange-
colored flashes indistinguishable from the flashes of Photinus castus
(?), which had been abundant at the first locality. Neither the latter
species nor the 5-flash species was observed at this second locality.

Thus in one evening in June at Washington one may encounter
pure colonies of five or more species of Photuris, and the vicinity of
Winona yields three species which occur at the same time but are
biologically very distinct, although, considered taxonomically, they
offer few reliable characters for recognition of cabinet specimens.
All these species have been until now commonly identified as pensyl-
vanica. If, however, the observer finds localities in which several of
these species are mixed, and their several females contribute to the
confusion of flashes, and if the observer collects but few samples
without noting their flashes, he is readily convinced that it is only
variation, and that there is no law of uniformity in the genus Photuris.

Variation in motive for flash, in the flash itself, as well as in size
and in pigmentation of body, must be admitted, and the writer is
far from satisfied on a great many points in this complex problem.
The female flash serving as a sex signal in response to a male flash
for the same purpose must be rarely visible to us. The flashes we
see from females must often be warnings or nervous responses to
irritation, but another suggestive phenomenon has been observed too
often to be ignored: Sometimes the familiar flashes of a small species
of Photinus male are observed excitedly courting a female, supposedly
of the same species, whose response flashes appear normal to its kind,
but when the electric light is thrown upon them one is startled to

10) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE., 17

find the intended bride of the Photinus is a large and very alert female
Photuris facing him with great interest. Does she lure him to serve
as her repast? Very often a dim steady light near the ground proves
under the flashlamp to be a small, recently killed male Photinus being
devoured by a nonluminous female Photuris, and females of the
largest form of Photuris (versicolor) have been found quietly feed-
ing on dimly glowing males of Photinus pyralis that had been wrapped
in silk in an orb web from which the spider had departed, the
Photuris female crawling on the web apparently in no danger of be-
coming entangled. Cannibalism has often been observed in captivity,
male Photuris being devoured by their supposed females; but the
writer’s observations and those of McDermott (1917), as well as
those of Williams (1917) and Hess (1920, p. 52), were made when
all our familiar Photuris were called by one name, and the sexes may
not have been conspecific. The accounts of Photuris pensylvanica by
all three of these writers seem to have been based upon two or more
species whose differences were interpreted merely as variation, but as
series of rapid flashes are referred to in each it appears that some
forms of the possibly composite species here called versicolor Fabricius
were included in the material for each of these studies.

Mistakes will be made by the most careful observer in his attempt
to record what he sees in connection with definite samples for sub-
sequent comparative study. Minor variations occur in the population
of a single species. Pure colonies are not often found. No satisfac-
tory timing device has been available. A watch producing half-second
ticks worn at the ear might offer sufficiently definite time rhythm for
more accurate estimates of flash duration and interval. The half-
second swing of a short pendulum on a stick held in the hand is suffi-
ciently accurate in spite of variation due to one’s irregular movements.
Its beat can be felt without looking away from the observed firefly,
and luminous paint on the apparatus has been found unnecessary.
After striking at a particular individual, two fireflies, perhaps of
different forms, may be found in the net, an unnoticed individual
having happened to be within the sweep of the net. Perhaps the
desired specimen is missed and an imposter receives the label of care-
ful observation, false when thus attached. But more often the trouble
of writing labels for single individuals in separate vials tempts one
to trust memory too far and vials become confused.

4 Some cheap watches tick four times to the second and if alternate ticks are
of different tone are very useful as a standard rhythm by which flash duration
and intervals can be estimated.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER II

Certain typical observations have been contrasted in the above ac-
count, and a short statement regarding measurement and interpreta-
tion must be made.

It should be obvious that since these phenomena are visible only
in the natural environment and represent the normal ways in which
undisturbed males seek to satisfy their mating instinct, laboratory
methods of exact measurement or controlled experiment are of no
use. It is difficult for one observer to contrast, verify, record, and
reconstruct all the factors of all the forms in this intricate problem,
even in the limited environment of Washington. Imagining the
ideal opportunity for observation, we might wish for two adjacent
pure colonies which could be observed and contrasted at leisure. In
any pure colony we must expect to observe (1) some variation in
the normal behavior of the seeking males, and (2) very irregular
behavior on the part of the females that have mated.

The courtship flashes of Photuris males appear to have become spe-
cialized in certain species from the normal short, single flash emitted
at rather regular intervals of 5 to 10 seconds, by increased frequency
in hebes (which flashes at 1- to 3-second intervals according to the
warmth or coolness of the evening), and in potomaca (which, on a
warm evening, may attain a rate of nearly two flashes per second), or
the duration of the light emission may be lengthened and interruptions
introduced as in the flicker-flash species versicolor, the protracted
tremulous flash of tremulans, the interrupted protracted coruscation
of the small marsh-inhabiting species pensylvanica, or the long
crescendo flash of lucicrescens.

The first-mentioned simple flash is given in such diverse colonies,
varying so in size, color, localities, and dates of appearance, that no
well-defined single species is discernible at this time, and no specific
name is here attached to samples. The frequency, pattern, and inten-
sity of the characteristic flashes of the males of several of the species
of Photuris described herein are diagrammed in figure TI.

NOMENCLATURE

Fears have been entertained that an unfamiliar name must be
adopted to replace Photuris. This name first appears in the 1833 edi-
tion of the Dejean Catalogue (p. 103), where 34 American species
are included, all but a few of which (perhaps all but three species)
are nomina nuda. Photuris versicolor Fabricius and hectica Fabri-
cius are valid species therein contained, and were it not for the query
after the latter name the designation of this species (hectica Fabri-

VOL)! 177,

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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No. Spectes
1 Primitive un-
named
2 cinctipennis
3 hebes
salinus
4 potomaca

5 frontalis

6 versicolor

7 versicolor var.

quadrifulgens

8 versicolor, triple

flash (Dela-
ware)

9 fairchildi

10 lucicrescens

Ir Tree-top spe-
cies, perhaps
the same as
lucicrescens

12 pensylvanica

13 pyralomimus

14 aureolucens

15 caerulucens

16 tremulans

Ila

Duration and
intensity of
single flash

Short; single

Moderate

Short; single

Weak

Short; single

Weak

Short; single

Bright

Short; single

Moderate

Composite of 3 to
5 or 6 very
short, bright
coruscations

Composite of 4
very short,
bright corusca-
tions

Composite of 3
very short,
bright, rapid
coruscations

2 short corusca-
tions separated
by an interval
Moderate

3 to 24 secs.

Very bright

Very short
and bright

Long; double
Moderate
¥ sec.
Moderate
Short
Weak
I sec.
Moderate
I sec.

Bright

FIREFLIES OF THE GENUS PHOTURIS—BARBER

EXPLANATION OF FIGURE I

Frequency
of flash

I each 5 secs.

I in 4 secs.

At I to 3 secs.

Rapid, up to 2
per sec.

At I-sec. or
less intervals

At about 5
secs.

do.

At 5 to 6 secs.

At 5 to 10
secs.
do.

do.
do.
I in 4 secs.
I in 4 secs.

I in 5 to 10
secs,

Color of light
Yellowish?

Yellowish?
Almost orange
Greenish
Greenish

Greenish

do.

Greenish

do.

do.
Yellowish?
Yellow
Bluish green

Greenish

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE Ty,

cius) as genotype by Motschulsky (1853) would demand recognition.
But Laporte’s revision (1833) of the genus Lampyris proposes a
different name, Telephoroides, for six valid species, including pensyl-
vanica, and LeConte’s 1852 rejection of “this uncouth name” claim-
ing that it was printed “as a French word” appears to be an unwar-
ranted action. LeConte’s apparent belief that a generic name is
invalid unless accompanied by diagnoses also led him to refer to
Photuris as “the hitherto unpublished name of Dejean.” In further
subdivision of the group, Motschulsky (1853) adopted both of the
above generic names and proposed seven new ones, designating geno-
types for all. Lacordaire (1857, p. 338, footnote 1) supports Le-
Conte’s attitude and rejects Motschulsky’s work, but in spite of the
latter’s designation of occidentalis Olivier as genotype of Telepho-
roides Laporte he credits this genus to Motschulsky and (p. 339,
footnote 5) designates pensylvanica DeGeer, with versicolor Fabri-
cius mentioned as synonym, as genotype. Gorham (1880) follows
Lacordaire but designates pensylvanica as the type of Photuris
LeConte. E. Olivier (1886) also ignores Motschulsky’s genotype
designations but rejects only five of his genera. In his 1907 work
E. Olivier does not allude to genotype and suppresses all nine
genonyms (credited to Motschulsky) under Photuris LeConte, but
in I910 the same author recognizes three genera, again ignores geno-
type designation, and arbitrarily lists the generic synonyms.

A future study must extricate the tangled nomenclature, but for
the present it is enough to claim that Motschulsky’s designation of
hectica Fabricius as type of Photuris Dejean is invalid under the
second paragraph of Article 30e of the International Code, and since
no other genotype designation is known the writer hereby designates
Lampyris versicolor Fabricius type of Photuris Dejean.

The genotype of Telephoroides, Lampyris occidentalis Olivier,
1790, designated by Motschulsky (1853), is unknown to me, and
Lacordaire’s designation of pensylvanica is invalid; but since the
former is cataloged in the genus Photinus by E. Olivier, 1910, our
continued use of the name Photuris for our North American species
may be justified even though the actual publication of the Dejean
Catalogue dated 1833 may be subsequent to the Laporte revision,
which appeared the same year.

A still more exasperating case is that of Pyractomena, in which
varied applications and spellings of the name have been incompletely
cataloged without application of the genotype principle. Revision of
all usages of the name is required, and we may even be forced to sup-

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 15

press its earliest valid form as a synonym of Photuris, but the proposal
of substitute names is unwise until more complete bibliographical re-
search is done and a better consideration of systematic relationships
is possible. It now appears that Lecontea E. Olivier, 1899, is the
proper genonym for the North American forms, although it differs
in only one letter from the pythid genus Lecontia Champion, 1880.

Those seeking to apply the law of priority and the genotype prin-
ciple to lampyrid genera may find the following chronological outline
suggestive :

1833. Dejean (p. 102) groups I1 species into a genus that first carries the name
“Pyractomena Dejean,” but, although specific names are listed from Klug,
Mannerheim, Latreille, and Dejean, no description of any of these species
by these authors has been found. Since all appear to be nomina nuda the
writer believes Pyractomena must be considered a nomen nudum of this
date, although he also believes that the citation of Dejean by authors
subsequently adopting his proposed genonym demands (article 19) that
evident lapsus calami or typographical errors be corrected. One of the
included species, marginata Latrielle, may be found to be valid if a
mention of marginata Linnaeus or Fabricius or Olivier can be found in
Latreille’s publication, but his only mention of this species that the writer
has found (Humboldt and Bonpland, vol. 1, p. 348, 1811) is casual. He
uses the French spelling without citation of author, and in the abbreviated
German translation of this paper (Germar Mag., vol. 1, part 2, p. 122)
the Latin name replaces the French form but without citation of Linnaeus.

1837. Dejean (p. 115) same as in 1833.

1843. Sturm (p. 76) in cataloging his collection adopts “Pyractomena Dej.,”
listing eight forms, all apparently nomina nuda, except the third species,
marginata, which is accompanied by citations to Linnaeus, Fabricius,
and Olivier. The generic name is therefore valid, with marginata Lin-
naeus, 1767, as its type, but this species is cataloged by E. Olivier, 1910,
as a Brazilian species of Photinus with only two references, the original
description and the redescription with figure by Olivier, 1790. This latter
figure looks so much like a Photuris that Pyractomena Sturm may be
one of its synonyms or subgenera, but until Linnaeus’ and Olivier’s types
can be identified with adequate modern specimens no certainty can be
felt that the figure represents the Linnaeus species.

1845. Melsheimer (Proc. Acad. Nat. Sci. Philadelphia, vol. 2, p. 304) described
two Pennsylvanian species using the genonym Pyratomena (c omitted),
but since he cites “Dej. Catal.” for the name, “a lapsus calami or a
typographical error is evident,” and the generic name must be considered
a homonym of that used by Sturm, but with lucifera Melsheimer, 1845,
as its type.

1847. Erichson (Wiegemann’s Archiv fiir Naturg.) adopted Pyractomena, cit-
ing Dejean, for a new Peruvian species, interrupta, which became his
monobasic type and is cataloged by E. Olivier, 1910, in Photinus, al-
though its bifid claws are more suggestive of certain groups of Photuris.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

1849. Solier (in Gay, Hist. Chile, vol. 4, p. 445) cites Dejean and thought he
adopted his invalid generic name, but spelled it Pyractonema (transposing
the » and m), for nine new Chilean species which have since stood as a
distinct genus under this name. His first species, compressicorne, is fig-
ured and is here designated genotype, but, as above argued, correction
of spelling is required and the name becomes a homonym. The proposal
of a new name is postponed pending a better knowledge of the limits of the
genus Lucidota, of which Pyractonema Solier appears to be a part.

1849. LeConte (in White’s Statistics of Georgia, p. 31, supplement) includes
no valid species.

1850. LeConte (in Agassiz, Lake Superior, p. 228) lists Lampyris borealis
Randall under Pyractomena Dejean, this being the basis of the below-
cited remarks by McDermott, 1917.

1852. LeConte (Proc. Acad. Nat. Sci. Philadelphia, vol. 5, p. 336) includes
five species under Pyractomena Dejean, borealis Randall being the fifth
species. A generic diagnosis being given, many authors have held this
as the first valid publication of the name.

1853. Motschulsky (Etud. Ent., 1852, p. 37) uses an e instead of an a in “Pyrec-
tomena Dejean” for which he designates “Pyractomena vitticollis Man-
nerheim” of Santo Domingo as genotype, but since this species appears
previously undescribed, although originally included (nomen nudum)
by Dejean, the generic description is held to be the first validation of the
specific name.

1857. Lacordaire (Gen. Coleopt. vol. 4, p. 321) suppresses Pyrectomena
(Dejean) LeConte as synonym of Photinus but later (p. 324, footnote 5)
applies it to one of the subgeneric groups, containing six species.

1880. Gorham (Trans. Ent. Soc. London, 1880, p. 32) treats Pyrectomena
(Dejean) Motschulsky, LeConte, citing vitticollis as type and recognizing
six species.

1899. E. Olivier (Bull. Mus. Hist. Nat. Paris, vol. 5, p. 371), not knowing
of the use of Lecontia Champion, 1889, for a genus of Pythidae, pro-
posed Lecontea as a new name for Pyractomena LeConte, 1851 (1852)
(into which he merged Pyrectomena Motschulsky, 1852) on the ground
that Pyractonema Solier, 1849, has priority. Lecontea E. Olivier is there-
fore isogenotypic with LeConte’s genus.

1917. McDermott (Can. Ent., vol. 49, p. 53) adopted the present writer’s opinion
(now reversed) and, holding the Solier and LeConte genonyms not
homonyms, designates Lampyris borealis Randall type of the latter.

From these facts it appears necessary to regard Pyractomena
Sturm as a possible subgenus or relative of Photuris and to discon-
tinue the use of the former name in the sense so long accepted.

A much more perplexing case also demands consideration but
seems to affect only the indexing of synonyms. To state that Pyrecto-
soma Motschulsky, 1854 (p. 39) is an isogenotypic synonym of
Photuris Dejean when its description was apparently drawn from a
species of Lecontea (Pyractomena) cannot but offend those who re-
gard genera as groups of species displaying the diagnosed character-

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 17

istics; yet the fact remains that versicolor Fabricius was originally
designated as its genotype, and Motschulsky’s subsequent “correc-
tions” (1855, p. 72) after seeing Fabricius’ type of versicolor can-
not change its generic nomenclatorial status. But to catalog his
taxonomic opinions it is still necessary to list Pyrectosoma versicolor
Motschulsky, 1853, 1854, and 1855 [not Fabricius] in the synonymy
of Lecontea, indicating that it is a pure primary homonym of the
synonym of Photuris and nomenclatorially not available for use as the
name of any species. In Opinion 14, the International Commission on
Zoological Nomenclature, 1910, has considered most of the principles
involved in this case, and in Opinion 65, 1914, a hypothetical case
almost identical in principle is treated, but these deal only with the
question of availability of the names.

These genonyms and genotypes, excluding the Pyractomena series
already discussed, may be listed:

Photuris Dejean, 1833, p. 103.
hectica Fabricius, genotype designated by Motschulsky, 1853, is not avail-
able because doubtfully included by Dejean (Article 30e of International
Code).
versicolor Fabricius, type by present designation. (This species is also the
originally designated genotype of Pyrectosoma Motschulsky, 1853.)
Photuris LeConte, 1852, p. 337.
pensylvanica DeGeer, designated by Gorham, 1880 (species not originally
included in Dejean).
Telephoroides Laporte, 1833, pp. 127 and 144.
occidentalis Olivier designated genotype by Motschulsky, 1853, p. 55 (cata-
loged in Photinus by E. Olivier, 1910).
pensylvanica DeGeer (versicolor Fabricius), genotype designation by Lacor-
daire, 1857, p. 339, footnote 5, is invalid because subsequent to that by
Motschulsky.
Pyrectosoma Motschulsky, 1853, p. 38.
versicolor Fabricius, genotype by original designation (therefore isogeno-
typic with Photuris Dejean, but characterization was drawn from mis-
determined specimens supposed to belong in Pyractomena of LeConte,
Lecontea Olivier).

Generic characters of Photuris are amply diagnosed by LeConte,
1852, but our more recent papers on fireflies appear to have con-
sidered the lunate last joint of the labial palpi and the cleft external
claw of all tarsi as unworthy of notice. In habitus all Photuris in
our fauna differ from other genera of fireflies in their more oval and
much less depressed form, which permits their sturdy, agile move-
ments to be so characteristically distinct. Supporting these peculiari-
ties in adults, their larvae are of such distinctive form and are so
adapted to free movement upon the surface of the soil that E. Olivier’s

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Li7

1907 and 1910 elevation of the group to subfamily rank distinct from
the Luciolinae is readily acceptable.

SPECIES OF PHOTURIS

The easy taxonomy of previous studies is reflected by Leng (1920)
in cataloging only three species of North American Photuris and
listing five supposed synonyms. Only two mild protests against this
simple concept of our forms are known to me, Wenzel (1896) hav-
ing remarked on “two forms of Photuris frontalis’ taken by him
at Anglesea, N. J., and Blatchley (1924) having sought to recognize
lineaticollis LeConte, 1852 (name omitted in LeConte, 1881), as a
distinguishable variety. Whether the better-described Telephoroides
lineaticollis Motschulsky, 1854, is identical with the form to which
LeConte had previously applied the name is immaterial at present,
and since the writer does not know the LeConte type of this species
the name is tentatively applied in the following table to a conspicuous
southern form displaying the character originally stated.

Two of the other species named by LeConte, 1852, congener and
frontalis, were confused by that author prior to his 1881 revision
and have ever since been misdetermined in all collections, the latter
name always being applied to the former species, and frontalis proper
being unrepresented by specimens. But in the Leng list congener
appears erroneously placed as synonym of divisa.

Of the four much older names hitherto considered conspecific with
pensylvanica, marginata Panzer, 1789, type locality “America meridi-
onale,”’ may be deleted from our lists since it is almost certainly a
South American species of Photuris not identical with any form in
our fauna, but since Panzer’s name is preoccupied by Lampyris mar-
ginata Linnaeus, 1767,° some other name must be used for his species
when it is reidentified. Photuris versicolor Fabricius, 1798, is not
a synonym of pensylvanica and must be recognized as one of our
species, but the writer’s observations would indicate a need of much
more critical study than has here been possible since his notes record
different behavior at different stations. The brief description of
vittigera by Motschulsky, 1854, appears applicable to the majority of

5 The source of Professor Brunniche’s sample which Linnaeus described can-
not be known and the type locality is America. This habitat was restated as
southern America by G. A. Olivier, 1790, whose figure shows long legs and
antennae suggesting Photuris, and was further restricted to Brazil by E. Olivier,
1910, who listed the species in Photinus. But as stated elsewhere in this paper
(see p. 15) marginata Linnaeus is genotype of Pyractomena Sturm.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 19

individuals inhabiting local fresh-water tide marshes, but this little
species is believed to be the original pensylvanica DeGeer, 1774, and
seems not to be elsewhere treated in literature unless it is one of the
two forms mentioned by Wenzel (1896) as frontalis.

SPECIES HERE DISTINGUISHED

Unsatisfactory as are many of the distinctions used in the accom-
panying table and in the appended comments, it is believed best thus
to emphasize the inadequacy of preserved specimens for specific
identification. The variation of characters customarily used for taxo-
nomic distinction is so obvious in the large series before me that,
had the specimens not been carefully collected to represent species
distinguishable on behavioristic peculiarities, no attempt at division
would have been made. Failure of such species to exhibit sufficiently
well-marked differential characters is probably not an uncommon phe-
nomenon, but owing to the existing dominance of taxonomy over
biology such species are too frequently ignored. A number of other
species of Photuris, believed to be new, are before me, but the for-
mality of naming them without the support of a definite knowledge
of their habits would be objectionable. It is believed that many more
biological units must be recognized and that many observers must
contribute opposing opinions before an agreement as to method of
taxonomic treatment is possible; but the long-accepted simplicity of
this genus is an example of our ignorance of one of the commonest,
most conspicuous, and supposedly best-known groups of insects.

PHOTURIS MALE GENITALIA

Since no specific distinctions in the male genital structures have
been observed, although abundant prepared material has been ex-
amined, we must give added emphasis to the supposed specific bar-
riers indicated in the courtship behavior, the ecological adaptations,
and the nuptial seasons of the different forms.

The male genitalia are unlike those of other lampyrids I have ex-
amined, as well as the four genera considered by Sharp and Muir,
1912, in that the sides of the “basal piece” are produced into long,
slender, clubbed, lateral processes extending beyond the apex of a
slender median lobe. A well-developed but very slender flagellum
or internal sac, often 4 mm. in length, armed with minute, flattened,
spinelike scales, is invaginated from the median orifice through the
median foramen and extends well into the coiled tube (stenazygos),
which passes through the basal orifice of the aedeagus and attaches

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

to the base of the median lobe. This flagellum appears capable of
being evaginated and probably is inserted to the spermatheca during
copulation; but except in length no distinctions were observed in
the different forms of which preparations were made. In no pre-
pared material has this flagellum been evaginated, and no duct at-
tached to its apex (probably the functional orifice) has been seen.

[The sketches (figs. 2, 3) are of the aedeagus of Photuris luci-
crescens from specimens taken in Delaware, Mr. Barber’s drawings
from Photuris frontalis not having been found. Figure 2 shows
dorsal, ventral, and lateral views of a cleared specimen, indicating
some of the internal structure, and figure 3 the dorsal and lateral views
on a larger scale of another specimen with the lateral processes spread.
The lateral lobes fuse with the dorsal surface of the median lobe
at about basal third, and are armed internally opposite this point
with a strong transverse ridge, which is sharply angulate at inner
third —McD.]

TABLE OF SPECIES OF PHOTURIS

1. Pronotum with or without infuscate area but always without a pair of
aval; red) discal: SHOtS. <i. wv os soaps ersten seine alae Ss a ale nitty laih ais, See eee 2

Infuscate median vitta of pronotum narrowed (sometimes interrupted)

in middle third by two conspicuous oval red or orange spots (pen-

sylvanica group comprising numerous similar species, of vaguely

dissimilar habitus but distinctive habits and habitats)............... 6
2 Pronotumpentirely ipale) yellowsmaicn. cease elonnis heen kal see Gee 3
Pronottim’ witheaiscal nluscatiOM\ynacitetalereiere eter oreneiete ie eyes iets aera 4

3. Elytra entirely black; metasternum concolorous with the yellow head
and thoracic sclerites above and below; fourth visible sternite with
apical margin pale, the lutescent area broad at middle, narrower
toward but not reaching the sides; apical infuscation of femora grad-
ual and hardly noticeable, but knees, including base of tibiae, pale;
length 8.4-11.5 mm. Type locality, Alpine, Tex. (flavicollis Fall, 1927,
not HE: Olivier; 1686): <).<)\ek. «sae oe 1. brunnipennis var. falli, new name

Elytra black with narrow sutural and broader lateral yellow margins
which are not continuous around apex; head, prothorax and meso-
thorax yellow above and below; the metasternum piceus; coxae and
basal five-sixths of femora yellow, the knees, tibiae, tarsi, antennae,
and four abdominal sternites black; length 11 mm. Type locality,
Raradisewisey. Place thenmatercre 2. brunnipennis var. floridana, new var.

4, /eronotal intuscate area median: front GlaVOUs. sors cvs sna.¢ auilemiee cee 5

Pronotal infuscation longitudinally divided by narrow median pale line;
front infuscate; emargination of penultimate sternite, size, sculpture,
etc., as in floridana except front broader in male, more than twice as
wide as one eye in same aspect. Type locality, “Missouri Territory.”

3. divisa LeConte

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 21

1177.

Fic. 2—Aedeagus of Photuris lucicrescens Barber. Cleared specimen showing
part of the internal structure. a, dorsal view; b, ventral; c, lateral.

1/7.

Fic. 3.—Aedeagus of Photuris lucicrescens Barber. Specimen showing lateral
processes spread. a, dorsal view; J, lateral.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

5. Pronotal infuscate area large, oval, not constricted and shading im-

10.

perceptibly into yellowish border; size, color, sculpture, and front as
in floridana but emargination of second luminous sternite less deep
and more broadly arcuate. Georgia (type locality), Florida, and
MORAG wich ecthdve cel se None Ee «hee Pee tone nec 4. congener LeConte
Pronotal infuscate area abruptly limited, usually twice as long as wide
and constricted at basal third; size larger (12 to 14 mm.), more
robust; coloration similar to congener but infuscation darker, pale
elytral margins continuous around apex, and femoral infuscation
reaching middle; front narrower, not wider than radius of eye.
Georgia (type locality), Maryland. Males emit short, slightly yel-

lowish flashes at less than 1-second intervals........ 5. frontalis LeConte
COXA HIMEGSCALE? cits ies vine 6 Mloteteretee sacs crete ene Tei iseketa:c) sl cisks ole \areleecten Tees F]
Coxae pale (except posterior pair in lineaticollis)..........0...0eeees 14
Size darger (10 to: £7 mim.) bi. ie ae amen ial wees sine) Ao terere Siar nee 8

Size small (8 to 10 mm.), pronotal infuscation usually broad with
smaller pair of reddish spots; color variable, the elytra usually with
short or moderate oblique pale vittae which are sometimes absent,
sometimes subentire, or the usually well-marked elytral infuscation
occasionally reduced to basal region by the increase of marginal and
sutural lutescence toward apex; femora infuscate in apical third;
tarsal joints pale with short apical infuscation; fourth visible sternite
infuscate, sometimes with posterior margin narrowly pale. Inhabits
fresh-water tidal marshes of Chesapeake estuaries, June and July;
males fly at top of marsh vegetation, poising to emit a protracted
double flash of greenish-white color, the first part very short and
immediately followed by a longer light emission lasting I to 2 seconds,
while making slight dip and rise; females remain deep in vegetation
CGlottigera: Motschuisky))....-:.0.2000-05 es0eiees 6. pensylvanica DeGeer

Fourth visible ventral segment mostly black, usually with narrow white
posterior margin (more variable in fairchildi) ; knees and usually
basal third of femora, as well as pale elytral markings, ochreous..... 9

Fourth visible sternite mostly white, the basal margin narrowly in-
fuscate, broadly so laterally; femora cream white with ante-apical
infuscation; first joint of hind tarsi white with apical fifth infuscate ;
length 12 to 14 mm. Abundant late in June in thick woods on alluvial
banks of Potomac River above Washington, D. C., the males emitting
short, greenish-white, very bright flashes at intervals of about 1 sec-
ond while flying through foliage................ 7. potomaca, new species

Infuscation darker; oblique elytral vitta usually shorter; male corus-
cations flickering or composed of three or more quick flashes. Chesa-
peake region and Minnesota, in latter region paler in color.......... 10

Infuscation more brownish; elytral vitta usually longer; male corusca-
tions, single flashes except in fairchildi which emits a double flash;
Minnesota, New York, Nova Scotia, and Virginia......:......ce0 Il

Elytral vitta usually well-marked but short; tarsal and antennal joints
strongly flavous basally; male coruscations of several types, three,
four, or several quick flashes differing in locality and brood, per-
haps indicating distinct forms. District of Columbia, Maryland, Dela-
WarevandMannesotassciac: eee eee ese 8. 2versicolor Fabricius

NO.

If.

13)

14.

10.

I FIREFLIES OF THE GENUS PHOTURIS—BARBER 23

Elytral vitta obsolescent; tarsal and antennal joints almost wholly
black; male coruscations consisting of four slow flashes. Cape
Pventyei Vial SUR tae 9. (versicolor?) quadrifulgens, new var.

Form slightly more robust; hind tarsal and antennal joints a little
more slender, the fourth to eighth inclusive measuring 44 mm.; sub-
humeral pale band usually broader than epipleural infuscation; length
12 to 13 mm.; males flying slowly at top of tall grass and over lawns,
dipping and flashing at about 5-second intervals, much like Photinus
pyralis but beginning its flash on downward flight. Selkirk, N. Y.,

TCL ee ed 2: ae A Oana eee 10. pyralomimus, new species
Form slightly more slender ; hind tarsal joints a little broader; antennal
joints shorter, the fourth to eighth inclusive measuring 3} or 3mm... 12
Antennal joints 4 to 8 inclusive measuring about 35 mm.; male corus-
cations single; habitat near Winona, Minn..................20c008- 13
Antennal joints 4 to 8 inclusive measuring about 3 mm.; male corusca-
tions double; habitat Cape Breton Island....... 11. fairchildi, new species

Generally similar in appearance to fairchildi, differing chiefly in having
shorter and narrower elytral vittae and somewhat darker coloration.
The antennae and posterior legs are proportionately somewhat longer,
the elytra a little wider, and the pronotum longer relatively to
the width than in fairchildi; the characteristic flash of the male is
unique, a I-second long, vibrating, tremulous coruscation. Habitat,
low land below Black Pond, Va.........-.-.. Ila. tremulans, new species

Size of pyralomimus (about 13 mm.) ; males emitting a slow, bluish-
green flash of about 1-second duration. Winona, Minn.

12. caerulucens, new species

Size slightly smaller (about 12 mm.;*abnormals measuring 10.5 mm.
and 13 mm.) ; males emitting a cae yellowish flash. Near Winona,

ING sp eae es Uma eGeLeee aanCaUAeMOnIGGr 13. aureolucens, new species
Elytra with well-developed oblique vitta; infuscation pale brown....... 15
Oblique elytral vitta obsolete, infuscation very dark................++ 17
Size small (10 to 12 mm.) ; labrum entirely pale or infuscate......... 16

Size larger (about 15 mm.) ; labrum pale at base, black at apex; more
robust, pronotal infuscation normal, oblique elytral vitta long; males
flying in abundance in July in swampy woods, poising in flight to
emit a long crescendo flash of greenish-white light of from 1 to 23
seconds’ duration, and of sufficient brilliance to illuminate foliage
several feet distant. Type locality, Priest’s Bridge, Patuxent River,
IMIG stance eas Seen Bie Se ee corto c 14. lucicrescens, new species

Labrum wholly pale (rarely slightly clouded) ; antennae long (7 to
8 mm.), slender; eyes smaller (2.0 to 2.2 mm. across) ; median pro-
notal infuscation very narrow, often interrupted at middle; oblique
elytral vitta sometimes short, rarely evanescent; males flying in
abundance about bushes in July, emitting short, rather feeble, slightly
orange flashes at about 1-second intervals. Inhabits willow-covered
fresh-water lowlands. Type locality, outlet of Black Pond, Va.

15. hebes, new species

Labrum black; antennae shorter and stouter; eyes larger (about
2.5 mm. across); median pronotal black area broad; male flash

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

much like that of Aebes (supra); inhabits salt-water marshes along
Chesapeake Bay. Type locality, a Baccharis thicket on sand spit at
Sherwood Forest, 7 miles northwest of Annapolis, Md. (July 7 and 9,
2S ;.) siey cease ten dyelon Sralate: fskere- 6 2b eeas Pai om ora Ne rade 16. salinus, new species
17. Size small (11 to 12 mm.), pronotal vitta normal, coxae and legs white
except ante-apical infuscate cloud on inner edge of front femora and
often some apical infuscation on tibiae and tarsal joints; pale margin
of elytra continuous around apices; first four visible sternites with
pale hind margins. Type locality, Sherwood Forest, near Annapolis,
TUG ERG Ye ar rel Ree eee SIRE GA ret aie 17. cinctipennis, new species
Size large (15 to 17 mm.); pronotal vitta usually interrupted; basal
half of femora and the coxae, except posterior pair, ochreous, the
latter partly or wholly infuscate; 4 black sternites without pale border.
Habits unknown. Florida and Louisiana...... 18. ?lineaticollis LeConte?

1. PHOTURIS BRUNNIPENNIS var. FALLI, new name

Photuris flavicollis FALL, 1927, not Olivier, 1886.

This conspicuous form was named after the present paper was
virtually complete, and its practical identity with brunnipennis was
not suspected until a specimen from Alpine, Tex., the type locality,
was obtained from Mr. Schaeffer and compared (January 1927) with
the Cuban specimens mentioned under the following variety. Such
close relationship between two striking forms, one inhabiting a tropi-
cal swampy region, and the other almost the summit of the Continen-
tal Divide in western Texas, should be supported by intermediate
colonies. Fall’s original description mentions the triangulate labial
margin, which is obscured by regurgitated material in my unique
specimen, but the divergence in this structure between Cuban speci-
mens of brunnipennis and the type set of floridana, mentioned below,
is noteworthy.

2. PHOTURIS BRUNNIPENNIS FLORIDANA, new variety

Eleven males taken by the writer on February 19 and 23, 1919,
at Paradise Key (Royal Palm State Park) about 40 miles southwest
of Miami, and four specimens (two males, two females) labeled
Miami, Fla., March 1920, P. Laurent, received from George M.
Greene, differ from the Cuban form, brunnipennis J. DuVal, in that
the yellow margins of the elytra are much broader, the metasternum
is wholly piceous, and the fourth visible sternite of abdomen is
piceous, except, rarely, faintly paler at middle, but never with the
broad white posterior margin as in the Cuban samples. The latter
consist of a male and female from Cayamas and Habana, deter-
mined by E. Olivier in 1911, and by Leng and Mutchler in 1922, sup-

NO. I FIREFLIES OF THE GENUS PHOTURIS—-BARBER 25

ported by a series of 4 males and 12 females from Sanitago de las
Vegas, Cuba. A better knowledge of peculiarities of different colo-
nies of brunnipennis in Cuba may show these differences to be insig-
nificant. The writer failed to make notes on the behavior of the
specimens he collected but believes they were flying low in the dense
“hammock” forest at dusk and emitting short single flashes. The
type locality is Paradise Key.

Type and 14 paratypes, U.S.N.M. No. 61oo1.

In floridana the labium is small, oval, slightly infuscate, and rarely
shows more than a feeble median tooth, while in Cuban brunnipennis
the infuscation is darker, the integument stronger, and the anterior
margin prominently tridentate, the teeth being formed by four equal
emarginations.

3. PHOTURIS DIVISA LeConte, 1852

Twelve specimens in National collection, with data as follows:
Topeka, Kans. (Popenoe), four specimens, one of them dated
July 19; Riley County, Kans. (Popenoe), four specimens, June 1, 4,
and 19; Kansas (Snow), two specimens; Baldwin, Kans. (Brid-
well), one specimen, and Lincoln, Nebr., July, collected at electric
light, one specimen. The type locality is “Missouri Territory,” and
there are three specimens in the LeConte collection bearing green
discs, which, according to that author’s labeling system, indicate “Ne-
braska, etc.” All specimens have the third antennal joint longer than
the second, as noted by LeConte, and all are males. The manner of
flashing appears to be unrecorded.

4. PHOTURIS CONGENER LeConte, 1852

The type stands as the seventh specimen of the series labeled
frontalis LeConte in the LeConte collection, apparently where that
author placed it when preparing his 1881 synopsis. Type locality
is Georgia. Nineteen specimens in National collection from Florida
(Daytona, March 1907, P. Laurent, one specimen received from
George M. Greene; Haulover (near Allenhurst), March 10 and 14;
Crescent City, May 25, and Lake Harney, Hubbard and Schwarz,
thirteen examples) and Texas (Columbus, July 3, Schwarz, one
example, and, without definite locality, from Belfrage collection, four
examples). One female from Texas shows no pronotal infuscation
and one each from Crescent City and Lake Harney have this infusca-
tion broken into a narrow prescutellar spot and a broader spot over
the head, but in the other specimens it is a large, ill-defined, elongate,
discal infuscation. Habits unrecorded. Length 9 to 11 mm.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

5. PHOTURIS FRONTALIS LeConte, 1852

This species appears not to have been recognized since its descrip-
tion more than 75 years ago, all students having followed LeConte,
1881, in applying the name to another species, congener, from which it
differs conspicuously in its larger size and more robust form. It occurs
in abundance in ravines and along the foot of a wooded bluff facing
Breton Bay, 2 miles from Leonardtown, Md., in July, the males fly-
ing rapidly through the foliage but not going high among the
branches. They emit very regular, quick, bright flashes of yellowish
color at intervals of about three-fourths of a second, abruptly dis-
continuing the flashes when they alight on foliage. Only two females
have been seen in several evenings spent in watching this species,
one, which was glowing faintly, in the grasp of a large phalangid on
the ground and more than half eaten, July 8, 1923, and one which
produced a fine streak of light as it descended to alight upon a leaf
some 8 feet above the ground, July 4, 1927.

The species was observed abundantly in the locality on July 13,
1923, June 19, 1925, and July 4, 1927; but was sought vainly on
June 18, 1926, and June 7, 1927. One male was caught at Sherwood
Forest, near Annapolis, Md., July 13, 1927 (P. G. Russell), and
another on St. George Creek, in St. Marys County, Md., July 1,

1931.
6. PHOTURIS PENSYLVANICA (DeGeer, 1774)

?Telephoroides vittigera MorscHULSKY, 1854, p. 60.

If the original types can be studied the above synonymy may
need revision, but of the species observed and collected by the present
writer only one appears referable to either of the descriptions origi-
nally accompanying the above specific names. This is the diminutive
species appearing in great abundance over the tall grass of the
Potomac and Patuxent tide marshes. The majority of the specimens
fit Motschulsky’s description, but only a few have the brown tint of
the elytra confined to the base as described by DeGeer, who records
the size as equivalent to 10 mm. His figure (pl. 17, fig. 8) is 14 mm.
in length but other familiar species are equally enlarged. His indirect
quotation from Acrelius informs us that they particularly inhabit the
prairies of Pennsylvania all summer, flying and shining like thou-
sands of sparks. Information about Acrelius has since come to notice
in the very interesting comment by Jones (Ent. News, vol. 41, p.
305, 1930), and it appears most likely that the type locality is within
the present city of Wilmington, Del., the southern part of which was

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 27

until a few years ago a fresh-water marsh and might then have
been called prairie. Ecologically it must have been practically iden-
tical with the marshes near Washington, over which vast numbers
of this little firefly may be observed. Samples studied consist of
about 90 specimens preserved by the writer on numerous visits to
their restricted habitat, and supported by two specimens from the
collection of George M. Greene, labeled Riverton, N. J., June 17
and July 3, 1899, which may be regarded as practically topotypes.

In many of them the oblique, pale elytral vitta is obsolete or
evanescent and in some the confluent infuscate area is narrowed by
widening of the lateral and sutural pale borders. In mid-April, 1927,
larvae were found by means of their lights to be very abundant in
the drier part of the marsh near the Shaw Lily Ponds, Kenilworth,
D. C., and when taken indoors they prepared their cells, pupated,
and issued as adults within a few days. Six weeks later (June 1)
the first adults were seen in the same locality, and 10 days later they
had become very numerous. By mid-July the numbers were con-
siderably reduced.

This species first attracted my attention on June 24, 1924, in the
Patuxent River marsh at Hills Bridge, Md., 20 miles east of Wash-
ington, and since it occurs in pure colony unmixed with other species
of Photuris, and flies at about the height of one’s head as he wades
in the marsh grass, it offers a very convenient contrast with the
treetop-frequenting forms that have hitherto been identified as pensyl-
vanica. Its very distinct behavior is so striking that it is strange no
observers have described it. Its habits have been noted often by the
writer, both in the Patuxent and Potomac marshes, and the follow-
ing composite account may better represent the species than scattered
detailed records.

One arrives after sunset, intending to watch. The dusk is settling
down over the marsh, and no firefly lights have yet been seen. Then,
in the darker, eastern side of an isolated alder bush comes the first
flash. Inspection discloses a male of this species rapidly ascending
a stem from the now very dark interior of the bush. Numbers of
others are thus appearing, but as yet they do not take flight, colors
of foliage being still visible. They flash at intervals in the darker
places, each flash being a very short, bright explosion of light suc-
ceeded by an equally short and abrupt interruption, followed immedi-
ately by the protracted second flash, the whole lasting I to some 3
seconds. When dusk has sufficiently advanced they fly, and others
appear all over the marsh. While producing the light they poise in
one place, with only a slight fall and rise in height, or perhaps while

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

ascending they make a slight spiral movement. No females can be
found, except by accident, until their presence near the roots of the
grass is understood, when the explanation of the peculiar stationary
coruscation of the male manifests itself. The grass standing verti-
cal, the response flash of the female could not be seen by the male
unless he poised directly over her and waited long enough for her to
answer his signal. Can such a high degree of adaptation of courtship
behavior to the peculiar structure of marsh plant growth be other
than an indication of long-established specific distinction? As one
cannot thus imitate the male and cannot expect the female to respond
after the rude commotion made by one’s close approach, flashlight
mimics are abandoned. Females confined in a screen-covered pan on
the bow of the skiff, which has been placed in the stream, partly
concealed in the wildrice, apparently answer flashes of males, but
the latter are too distant and the cloud of mosquitoes, as well as the
belief that males recognize and avoid abnormal environment, dis-
courages perseverance.

7. PHOTURIS POTOMACA, new species

No other characters than those given in the key have been noticed,
and variation is found even in these. Five of the 24 males show
greater extent of the basal infuscation of the fourth visible sternite,
approaching the condition in versicolor. In two of the same series the
apical infuscation of the tarsal joint is diffused basally and in the
eight females taken with these males the fourth sternite is black or
only narrowly bordered with white. Abundant and conspicuous as
this species is in the shore woods of the Potomac above Washington,
the writer has failed to preserve an adequate series, as only two lots
are available: 19 males and 4 females from Offutt Island (type lo-
cality) in the Potomac, 2} miles below Great Falls, Md., June 23 and
24, 1926, and 9 specimens, 3 of them females, taken June 30, 1926,
on the Virginia shore at Stubblefield Falls, near Plummers Island,
Md. Another male was taken July 1, 1926, by the river near Black
Pond, Va. As already stated, the males emit very short, greenish
flashes at regular intervals of about 1 second, while flying in woods
along the river banks.

Type and 32 paratypes, U.S.N.M. No. 61002.

8. PHOTURIS VERSICOLOR (Fabricius), 1798

The type locality is recorded as “Habitat in America Borealis Dom.
Hirschell,” but a record of the residence of, or places visited by, Mr.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 29

Hirschell in this country has not been found by the writer. The useful
characters of the original description are: large size; black antennae
with the articles a little pale at base; black elytra with margin, suture,
and a short vitta yellow ; black legs with knees yellow.

About 100 recently collected specimens from the Chesapeake re-
gion display this combination, but, although they are not believed
to represent one homogeneous species, the writer’s notes on behavior
are insufficient for their separation. All notes refer to flickering
coruscations, but unfortunately no timing device other than the ex-
perience from timing photographic exposures was used in making
observations on any of them. The discrepancies in my records are,
however, too great to ascribe merely to varying judgment. Although
possibility of errors cannot be denied and temperature alters behavior
to some extent, the following notes on observed flashes are offered
as perhaps of help in future observations.

An early form was found in a field in Rock Creek Park (June 109,
1924, and June 2, 1925) and along the Virginia shore of the Potomac
River near Stubblefield Falls (June 20, 1924), males flying slowly
5 to 15 feet above ground, emitting a rapid series of five or six short
flashes of moderate intensity and greenish hue in less than one-half
second and at short intervals. Numerous males were attracted ex-
citedly to the mimic of the female light by a flashlight dimmed with
green leaves and fingers, while the writer stood concealed in foliage
at edge of field. Basal third to half of first joint of hind tarsi is
yellow in preserved samples, except in two specimens (June 2, 1925)
in which yellow extends to apical fourth. Elytral vitta varies from
short basal vestige to two-thirds entire.

In a field near Cabin John Postoffice, Md. (June 7, 1927), males
emitted three or four short flashes in about a second, followed by a
long rest, but when observed the temperature was falling rapidly
after a warm afternoon and we may suppose that persistent males
were acting abnormally.

In the field below Black Pond (10:30 p. m., August 2, 1927)
among the few females and very rare males of hebes and lucicrescens
then surviving, a single male versicolor(?) flew swiftly along the
edge of the woods, 15 to 30 feet above the ground, emitting greenish
flashes in series of four in about three-fourths of a second and at
6- to 8-second intervals, the fourth of each series being much less
brilliant than the first two. This individual was observed to fly 300
yards or more before descending within reach of the net, and since
it displays no characters by which it can be separated from the above

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

series it is regarded as a stray from the earlier brood, belated, per-
haps, by having transformed in a place chilled by a flow of cold
spring water. Its elytral vitta is a mere vestige, and the antennal joints
are not pale at base, but this may also be the result of a cold environ-
ment.

Among confusion of flashes by several species two specimens emit-
ting only three flashes were taken June 8, 1927, at Breton Bay near
Leonardtown, Md., one resting on foliage and leisurely producing
three flashes in about 14 seconds at rather long intervals, the other
flying and emitting three flashes in one-half second at about 5-second
intervals. In these the first joint of the hind tarsi is about three-
fourths yellow.

Mr. McDermott observed a form at Claymont, Del., on June 11,
1927, which flew 3 to Io feet above the grass, emitting three rapid
successively brighter flashes at 2- or 3-second intervals.

Among what seemed to be five species of Photuris active at the
mouth of a sharp ravine in Sherwood Forest on the Severn River
near Annapolis, Md., June 29, 1927, were a few swift-flying males
emitting a very rapid and brilliant flickering flash with perhaps eight
or more vibrations too fast to count, in about one-half a second, at
intervals of about 3 or 4 seconds, and at distances between flashes of
from Io to 20 feet. Attempts to distinguish the series of seven males
and five females preserved from this locality have failed.

Near Winona, Minn., July 6, 1926, a series of 13 males was pre-
served by Miss E. Myers and Mr. Boland, who noted that they flew
2 or 3 feet above the tall weeds, emitting usually five greenish flashes
as fast as one could count at intervals of perhaps 30 seconds. These
specimens average a little smaller in size and are paler in color but
otherwise appear not separable from the above forms.

9. PHOTURIS VERSICOLOR QUADRIFULGENS, new variety

Three specimens captured out of a score observed May 21, 1927,
near Cape Henry, Va., are darker colored, with scarcely a trace of
the basal paleness on antennal or tarsal joints, the elytral vitta wholly
absent in one specimen, an obsolescent vestige in another, and very
short in the third, and the elytral apices black in two specimens, while
the pale margin is very narrowly continued around apex in the third.
They were emitting greenish, bright, perhaps one-half-second flashes
in series of four, with short intervals of about a second and longer
intervals of a quarter to a half minute, but the evening was not cold,
and mosquitoes were very aggressive. The specimens were found

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 31

flying among scattered pines on the old sand dunes bordering a small

fresh-water marshy area near the south end of the bridge over Long

Creek about a mile east of Lynhaven Inlet. Length 13 to 14 mm.
Type and 2 paratypes, U.S.N.M. No. 61003.

10, PHOTURIS PYRALOMIMUS, new species

Size and habitus of versicolor but a little more robust and less
deeply infuscate. Individuals vary from pale brown with ill-defined
pale marks to dark brown with sharply defined yellow markings. The
lutescence of hind tarsal and antennal joints varies greatly, that of
the former occupying one-third to five-sixths of the first joint. In
about one-fifth of the specimens the epiplural infuscation is enlarged.
The species was observed by the writer in vast numbers July 3, 1924,
near Selkirk, N. Y., flying slowly about the lawns and hayfields, the
males dipping, flashing, and poising at tips of tall grass very much
like Photinus pyralis, but emitting their half-second flash during the
descent as well as the ascent. Thirty-three males and three females
preserved.

Type and 35 paratypes, U.S.N.M. No. 61004.

11, PHOTURIS FAIRCHILDI, new species

Varies from pale elytra with basal infuscation (three specimens)
through darker shades of brown on infuscate areas of elytra to the
normal dark-brown infuscation (two specimens) more common in
the genus. Fourth visible sternite is narrowly bordered with white in
three specimens, the white more extended in others, until in three
specimens the infuscation is only conspicuous at sides. The slightly
shorter antennal joints and the uniform size of about 12 mm. are
practically the only differences observed to support the distinct be-
havior and remote habitat. Ten specimens, one a female, were received
from Graham Fairchild, with the information that they were caught
over marshy ground at Baddeck, Nova Scotia (Cape Breton Island),
about 9:30 p.m. on July 14, 1927; that they fly rapidly and emit two
medium flashes separated by an interval about twice as long as one
flash, but that the flashing is not very regular.

Type and 9g paratypes, U.S.N.M. No. 61005.

11a, PHOTURIS TREMULANS, new species

This species has been taken in low ground below Black Pond, Va.
It resembles the type specimen of Photuris fairchildi but is somewhat

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

darker and has short and narrow elytral vittae ; the antennae and pos-
terior legs are proportionately longer, the elytra somewhat wider, and
the pronotum tends to be longer relative to the width. The very char-
acteristic male flash, a long tremulous coruscation lasting one-half
second to a second, differentiates this species clearly from others of
similar appearance.

Type and 4 paratypes, U.S.N.M. No. 61006.

12, PHOTURIS CAERULUCENS, new species

Form and colors as in the paler variety of versicolor from vicinity
of Winona and hardly distinguishable from it in the cabinet. The
shorter antennal joints, slightly broader first joint of hind tarsi, and
very slightly smaller average size help in the recognition of preserved
specimens of the present species, whose lights were observed as very
different from the versicolor also present there. According to the col-
lectors, who called this species the “slow blue,” the normal male flash
is a steady bluish-green light of about a second’s duration, dimly visible
for some time after the flash. Twenty-six males and four females
were collected by Miss E. Myers and Mr. Boland on July 8, 1926,
over damp ground close to a tamarack swamp near Bluff Siding (type
locality) in Wisconsin, 10 miles east of Winona, Minn., in company
with another species (aureolucens), and four males and two females
were preserved two days earlier near Stockton, Minn. (10 miles east
of Winona), where they were less abundant among the pale variety of
versicolor.

Type and 35 paratypes, U.S.N.M. No. 61007.

13. PHOTURIS AUREOLUCENS, new species

Form and coloration of caerulucens, from which it is almost indis-
tinguishable in cabinet specimens. The smaller size and slightly more
slender antennal and hind tarsal joints are inadequate recognition
marks, but the information kindly supplied by the collectors states
that this species emits a single, short yellowish flash not to be dis-
tinguished from that of Photinus castus, and flies about the tops of
tall weeds in marshy ground, appearing in the dusk before caerulucens,
with which it occurred but from which it is conspicuously different
in the color and the duration of the flash. Twenty-three males and
one female collected near a tamarack swamp in Wisconsin, near Bluff
Siding, 10 miles east of Winona, Minn., July 8, 1926, by Miss E,
Myers and Mr. B. Boland.

Type and 23 paratypes, U.S.NM. No. 61008,

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 33

14. PHOTURIS LUCICRESCENS, new species

This is the largest, palest-colored, and most brilliantly luminous of
the species encountered by the writer in the Chesapeake region, but
much remains to be learned of its behavior. It may be the species
figured by G. A. Olivier, 1790, as pensylvanicus DeGeer. Cabinet spec-
imens may be recognized by their pale coxae, brownish color of in-
fuscate areas, usually strong development of the lutescent borders and
oblique vitta of elytra, and the irregularly lutescent areas in the first
three or four visible sternites. The series from the densely wooded
low shores of the Patuxent River at Priest’s Bridge, Md., 20 miles
east and slightly north of Washington, D. C., has been chosen as
typical because in this locality no other species was observed during
July, and especially because the puzzling short flashes in the treetops,
mentioned below, appeared to be absent. Here the myriads of flashing
males usually flew lower in the forest, and emitted lights of greenish-
white color, which began dim, grew brighter, became very brilliant,
illuminating foliage for several feet around, and ended abruptly,
having lasted from about three-fourths second to 24 seconds, as timed
by a pendulum of one-half-second beat. The type, allotype, and 11
paratypes were taken from this colony on June 29, July 1, and July 22,
1927, and the behavior of the numerous population of the species was
also watched on the evenings of July 5 and 12. On the latter date
special attention was given to the presence with lucicrescens, in woods
of adjacent valleys, of a similar or identical form flying about the
upper branches of the trees and emitting extremely short (perhaps
one-tenth second) and bright flashes at intervals of 3 to 5 seconds.
Satisfactory samples of those thus flashing could not be obtained, but
on July 22 a male observed to be emitting these instantaneous flashes
was caught by a wind eddy and descended, still flashing, within reach,
where it was illuminated by the flashlight beam and taken. No char-
acters have yet been found by which it can be differentiated from
typical lucicrescens. Among a series of males from Sherwood Forest,
Severn River, near Annapolis, Md., July 5, three specimens were
thought to be giving these very short flashes, but not having been
illuminated by flashlight before netting it was feared that a nonflash-
ing lucicrescens might have been taken.

In some localities, or under some conditions, this species appears to
poise for its long flash. At other times and places what may be this
species flies a zigzag course over the bushes, coruscating only while
on a short sidewise flight at nearly right angles to the general direction
of its advance, and in some localities the size averages a little smaller

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: si7,

and the flash shorter. At times a definite vibration to the light can be
seen. F. A. McDermott, at Claymont, Del., July 19, 1927, describes
in a letter the strong crescendo flash as “unquestionably vibrating”
and recounts his attraction and capture of males by producing short
flashes with a small pocket flashlamp covered by two layers of plan-
tain leaves. The writer’s success in similar attempts has been variable
and leads to the belief that the searching males have extremely good
vision and readily perceive an enemy unless the observer stands con-
cealed in foliage. His most striking success, however, was not with
a flashlight but by the use of the light of the fireflies dying and glow-
ing brightly in the cyanide bottle. The latter was held concealed, its
light being exposed for very short periods by quickly opening and
closing the hands, and several males were observed to alter their
course and approach as if for courtship.

Although the dates on preserved specimens range from June 21 to
August 29, the period of chief abundance usually covers about 3
weeks in early July, after which males are less in evidence and through
August most of the individuals encountered are females. The 136
specimens are from the following localities: Maryland—Priest’s
Bridge (type locality), Plummers Island and vicinity, Lanham, Ber-
wyn, Sherwood Forest, and Breton Bay; Washington, D. C.; Vir-
ginia—Hunting Creek (1 mile south of Alexandria), Black Pond,
near Great Falls; Delaware—Claymont. (One specimen seen at
Louisville, Ky., June 1945.—McD.)

Type and 135 paratypes, U.S.N.M. No. 61009.

15. PHOTURIS HEBES, new species

Forty-two specimens are preserved from the type locality, Black
Pond, Va. (Potomac River, 2 miles below Great Falls), collected
June 28, 1925, July 21 and 26, 1926, July 3 and 28, and August 2,
1927. These are of small size (about 11 mm. long) and have rela-
tively long antennae, and pale (sometimes slightly infuscate) labrum,
but exhibit considerable variation in the extent of the elytral vitta,
which usually passes the middle, although it is sometimes evanescent,
or may be broader and attain the apical fourth of elytra. The pro-
notal infuscation is rarely interrupted by medium coalescence of the
orange spots, but may sometimes attain a width approaching that of
one of the orange spots. Fourteen specimens from Chalk Point (7
miles south of Annapolis, Md.) were preserved July 13, 1926, out
of many seen flying about Baccharis bushes and over the intervening
tall grass bordering the salt water. The flying males emitted short,

NOS £ FIREFLIES OF THE GENUS PHOTURIS—BARBER 35

sharp, yellow flashes at about I-second intervals, in strong contrast
with the much brighter, greenish flashes of another species oc-
cupying adjacent trees. On searching for the females they were found
in numbers in the bushes and grass. At this place the impression of
the yellowish character of the light was very strong, while in the
type locality the impression of contrast was less marked.

Four other specimens seem referable to this species and are from
Plummers Island, Md., July 9 and 24, 1902 (H. S. Barber), the Vir-
ginia shore near the same island, July 21, 1923 (H. S. Barber), and
Lakeland, Md., July 5, 1909 (F. Knab). Preserved samples of hebes
resemble the average specimen of pensylvanica in their small size and
dorsal coloration, but the intermediate joints of the antennae are
longer and the coxae are pallid. The very similar specimens found by
Wenzel in the sea-water meadows at Anglesea, N. J., have much
shorter antennal joints and are here referred to salinus; they are
probably one of the “two forms of Photuris frontalis”’ taken there
and mentioned by Wenzel, 1896. A closely related form inhabiting
the Florida Everglades is omitted, the writer having failed to make
sufficiently definite observations upon its habits.

Type and 59 paratypes, U.S.N.M. No. 61010.

16. PHOTURIS SALINUS, new species

Similar in size, form, and flashing habits to hebes but peculiar to
the drier margins of salt marshes near Chesapeake Bay, and differing
in having the labrum black, the antennae shorter and slightly stouter,
the eyes larger, and the infuscation of the mesopleurae more pro-
nounced.

Type locality, a Baccharis thicket on sand spit at Sherwood Forest,
7 miles northwest of Annapolis, Md. (July 7 and 9, 1928). Other lo-
calities: a Baccharis-bordered salt-grass area on St. George Creek,
St. Marys County, Md. (July 1, 1931).

Specimens doubtfully referred to this species were collected near
Lloyds, Dorchester County, Md., on July 10, 1907, by the writer, and
at Anglesea, N. J., by H. W. Wenzel, probably being one of the two
forms mentioned by him (1896) as Photuris frontalts.

Type and 43 paratypes, U.S.N.M. No. 61011.

17. PHOTURIS CINCTIPENNIS, new species

There is a possibility that the small (11 to 12 mm.) species for
which this name is proposed may be identical with either Photuris
lineaticollis LeConte, 1852, or Telephoroides lineaticollis Motschulsky,

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

1854. The small size, almost wholly white legs, white elytral epi-
pleura, deep black elytral disc, broad pale elytral margins, and usually
total absence of oblique median pale vitta make this form conspicu-
ously distinct in collections. Unfortunately its distinctness was not
recognized at time of collection, when attention was concentrated
upon other species, and its lights were not particularly noted. Among
the flashes observed on that occasion, and not ascribed specifically to
individual fireflies, were, however, only the more ordinary short and
regular flashes commonly given by restless females of most Photuris
species and by searching males of a few species. Two females were
collected at Breton Bay, Md., July 8 and 13, 1923, and a series of
three males and six females at Sherwood Forest (type locality) on
the Severn River near Annapolis, Md., June 28 and 29, and July 5,
1927, only females being taken on the last date.

Type and Io paratypes, U.S.N.M. No. 61012.

18. PHOTURIS LINEATICOLLIS LeConte, 1852
?Telephoroides lineaticollis MorscHuLsky, 1854.

Under this name are placed six very large, dark-colored female
specimens from Florida and Louisiana, as listed below. There is, how-
ever, considerable doubt about their identity, since the writer failed
to notice a specimen in the LeConte collection which might be the
type of that author’s short remark of 1852 validating the nomen nu-
dum of the Dejean Catalogues. The identification of lineaticollis
Motschulsky by Gorham, 1880 (p. 110), from Quebec, requires re-
examination. Blatchley (Can. Ent., vol. 56, p. 165, 1924) has quoted
this remark and added some discussion, but states the length as
14 mm., agreeing in this respect with the form described by Mot-
schulsky, 1854. In only two of the specimens, all of which are larger
than the length just stated, is the median infuscation of pronotum of
linear form, the other four having this dark line interrupted broadly
at middle, forming a larger rounded anterior spot and a narrow pre-
scutellar spot.

This is probably our largest North American firefly, and if the
large area of the urate reflector in the lumious segments is an indica-
tion of its light, it may be our brightest-flashing species as well. No
notes on its habits are now available, however. The six specimens
are labeled as from Archer, Fla., March 1882 (Koebele) ; Hillsboro
County, Fla., May (Hubbard and Schwarz); Lakeland, Fla., April
1912 (G. G. Ainsley) ; Duval County, Fla., and Covington, La.,
May 28 (Soltau).

ADDENDUM

NOTES ON SOME GENERAL CHARACTERS OF
NORTH AMERICAN PHOTURIS

By Frank A. McDermott

A somewhat detailed examination has been made of 28 specimens
representing 19 species and varieties of Photuris which Mr. Barber
had assembled as representing most of the species discussed in the
foregoing monograph, and also of type specimens of tremulans and
salinus. Measurements and points of particular difference or interest
are given later in this section.

Certain characters are in general very similar in all the species,
these being of some generic importance. There is, of course, some
variability between different specimens of the same species in all
characters ; such phenotypic differences are to be expected, and there
are instances where the variation may overlap between species, for
example in over-all length or width. It is difficult to describe ac-
curately in words, or even to illustrate properly, the shape of some of
the appendages—e.g., the labial palpi—though an attempt has been
made to make them recognizable. Some of these general features are
discussed in detail below, and in some instances may be compared
with the generic characters as given by LeConte, Olivier, and others.
For the sake of reference, the generic descriptions by LeConte, La-
cordaire, and Olivier are also given.

Pronotum.—Unlike the conditions in the commoner species of the
genera Photinus and Lecontea, the carapacelike pronotum does not
completely cover the head, so that, as viewed from above, a portion of
the eyes and frons is visible. The shape of this structure is generally
roughly scutate, or perhaps more accurately, rounded ogival, broader
than long, and with rounded angles at the posterior lateral corners.
In most species there is a median pigmented area, usually consisting
of a central dark-brown or black figure, between two orange or pink
areas ; in some species this pigmented area is absent, being represented
by a merely shaded or dusky spot in the otherwise uniform chitin.
Outside of this pigmented area, the remainder of the pronotum may
be opaque yellow or white, translucent, or even transparent. The
shape of the dark pigmentation is at least somewhat characteristic of
the species, though somewhat variable in different specimens. The

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

characteristic form, as represented by Ph. versicolor, is a T on a tri-
angular base, the apex of the latter coinciding with the median line
at the anterior edge of the pronotum, and the cross bar of the T lying
along the posterior edge; this T-form is subject to several specific
modifications, as given in the descriptions of the species. There may
or may not be a median sulcus in the pronotum, and there would seem
to be some doubt as to whether, when present, it is a natural character
or an artifact resulting from distortion in drying.

In the specimens examined the ratio of width to length of the pro-
notum varied from 1.2 to 1.5; no relation was evident between this
ratio and the over-all size of the insects. The proportion of the total
length (pronotum plus elytra) represented by the pronotum varied
from 18 to 21 percent, averaging about 19.7 percent.

Scutellum.—tThis small structure is roughly kite- or coffin-shaped,
and varies in coloration more or less, and to some extent in outline, with
the species ; again, it is difficult to express the exact shape in words.
Anteriorly to the scutellum, the two mesonotal plates may usually be
seen sufficiently to note the color, which is frequently the same as that
of the scutellum.

Elytra.—Since the elytra represent about 80 percent or more of the
total area of the insects as seen from above, differences in them are
the most easily recognized characters. In general, two types of elytral
outlines are found in these species: (1) Those in which there is no
marked widening or outward curvature of the lateral edges, and which
are therefore described as parallel or practically parallel; (2) those
in which such a widening is definitely noticeable, usually as the result
of the presence of a distinct margin, and where the resulting outline
is at least subparallel and approaches a long oval. This condition re-
sults in a considerable range of variation in the ratio of length to
width, the figures found for the species embraced here varying from
2'3 to.3'8t.

The base or ground color of the elytra varies from a very dark
brown—nearly black in some specimens—to a pale grayish tan. Per-
haps it is in the base color that the greatest amount of individual varia-
tion will be found. For instance, three specimens of Ph. lucicrescens
in Barber’s collection, dated 1927, are all light, although there is some
difference between them; two taken by the writer in Wilmington in
1948 are both much darker than Barber’s specimens, although other-
wise very similar. Of course the expression of such color tones in
words may convey to another reader a different shade from that in-
tended, and hence an attempt to give a very definite color classification

NOs E FIREFLIES OF THE GENUS PHOTURIS—BARBER 39

is not justified. Another difficulty is the darkening of the specimens
with age; the originally practically pure white of the luminous seg-
ments becomes eventually a brownish yellow, and other light areas
undergo a similar darkening; presumably the darker portions also
deepen in tone. Still a third factor is that in examination under a bin-
ocular microscope with intense illumination, all colors appear lighter
and brighter than under general illumination; hence the appearance
under the latter condition may really be more significant than under
the microscope.

In the majority of the species, a rather definite lighter border or
margin is present on both the lateral and sutural edges of the elytra,
and these margins may be continuous by meeting around the tips of the
elytra. In some, the lateral margins are relatively quite wide for a con-
siderable part of the elytral length, and are associated with an increase
in the maximal width. The sutural margin is usually rather narrow,
not much more than a line. A further feature characteristic of many
species is a light-colored stripe, called a vitta by LeConte and Barber,
beginning at or near the shoulder (humerus) and extending length-
wise of the elytron, and obliquely so as to approach the suture; this
stripe may vary in length from one-fourth to seven-eighths of the
elytral length, a variation of some diagnostic value. It is usually fairly
wide at the anterior end, narrowing rapidly at first, and then gradually
for the greater part of its length, eventually becoming indefinite and
no longer traceable. Usually each elytron has one or more ridges or
costae, which appear to be lines along which there is an exaggeration
of the general tuberculation. These costae usually begin at or near
the humerus, and may extend for almost the entire length of the ely-
tron, but more frequently end indefinitely at one-half to two-thirds
the length. They tend to diverge, and when oblique stripes or vittae
are present, the most prominent ridges may mark the middle of these
stripes. However, it has been noted that the number of such costae is
not necessarily constant in all specimens of a species, and although
not infrequently given in the description of a species, they appear to
be unreliable as a specific character. This is also true of the hair,
which is a prominent feature of most species; occasionally this may
be locally developed in a manner requiring mention, but usually it is
a general condition and rather variable.

Head.—Viewed from the front the head appears to be composed
mainly of the two large eyes and the frons—the area between the eyes
and bearing the antennal sockets. The width of the frons, its color,
unusual details of the antennal sockets, the slope or divergence of the

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

interocular margins, etc., may be of diagnostic importance, but for
most species the measurements do not provide means of identification.
The ratio of the frontal width across the eyes to the total length varies
from 0.18 to 0.24 for the specimens examined, without parallelism to
the total length. In these species, the frons is usually depressed
medially.

The terminal joint of the maxillary palpi is the portion of this
structure most easily observed; this joint is usually long-conoidal in
outline, flattened and lighter on the inner surface, and frequently the
tip is flattened or bent to give a finger-tip appearance ; usually this tip
is rounded, sometimes nearly straight across, and may appear as a
sharp point by lateral view. The labial palpi, described as lunate by
LeConte, is rarely even approximately crescentic in these species—at
least it is a very asymmetric crescent. Perhaps it is best described as
being of a long, narrow mitten shape, with the “thumb” projecting at
a right angle; this thumb may be curved at the end, and may have a
low protuberance at its base. For most of the species, the variations
in outline are slight.

The labrum (perhaps more properly the clypeus) shows some varia-
tion ; the edge may have one or more small projections, and the whole
structure may be short, not completely covering the closed mandibles.
The mandibles are curved, sickle-shaped rather than semicircular, and
under the microscope may appear to be hollow; they are brown, lighter
in the proximal portion, and although appearing rather thick (0.05-0.1
mm.) for insects of the size of these, are sharp-pointed.

Antennae.—Perhaps next to the elytra and the pronotum, the an-
tennae are the most conspicuous features in Photuris. They are longer
than in many lampyrids, but are simple, 11-jointed, and slightly taper-
ing. Their length, expressed as a fraction of the total length of the
insects, varies from 0.455 in pensylvanica to 0.69 in hebes, the ma-
jority being between 0.5 and 0.6. The third joint is little if any longer
than the second; the first joint is usually the longest, and any one of
joints 4 to 10 is usually as long as or longer than the sum of the
second and third; the eleventh is usually somewhat shorter.

Sternites——The first four visible sternites are of about the same
length, and usually mainly some shade of brown, the posterior one
frequently being one-third to one-half white; the posterior edges are
but little sinuate, being nearly straight. The sixth and seventh ster-
nites are completely white and represent the main luminous area; the
eighth is usually much smaller and white, but not apparently luminous,
and in a number of species it bears a long (0.25 mm.) median projec-

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 4I

tion, with a base which may be broad or narrow. The posterior edges
of the sixth and seventh sternites are usually more or less emarginate
or “notched,” sometimes deeply ; usually both are 1.3 to 1.5 times the
length of any of the first four sternites. The “foveae” (points of mus-
cle attachment), noticeable on the ventral side of the luminous seg-
ments in Photinus and Lecontea, are rarely observable.

The aedeagus was extruded in 10 of the 28 specimens examined ; in
all cases it was of exactly the same type as far as could be determined
without dissection ; it varied from 1.75 to 3.0 mm. in length, represent-
ing 17 to 21 percent of the total body length, and tending to be longer
in the larger species. The same type has been found in dissections of
Ph. versicolor and Ph. lucicrescens collected in Delaware by the writer,
and in Marthas Vineyard, Mass., by Dr. Frank M. Jones, and is very
similar to that of Ph. jamaicensis collected in Jamaica by Dr. John
B. Buck.

Legs.—The legs of Photuris are proportionately much longer than
those of Photinus and Lecontea, and in occasional specimens impress
one as being unusually long, especially the posterior pair. Measure-
ments show that these posterior legs vary in length from about 0.65 to
0.85 of the total length of the insects, averaging about 0.75. The outer
claws are bifid on all legs, and sometimes there is a small protuberance
at the base of one or both claws. Pronounced tibial spurs are present
on the two posterior pairs of legs. Claws and spurs are usually a
clear brown. The lobes of the fourth tarsal joint usually extend well
toward the claws, covering most of the fifth joint. Each lobe has a
furry pad on the under surface, which may be gray or black, instead
of yellow or brown.

The generic descriptions referred to above are given here.

LeConte, J. L. Proc. Acad. Nat. Sci. Philadelphia, vol. 5, pp. 331-
347, 1852. Photuris Dejean, p. 337:

Antennae I1-jointed, slender, elongated, joints 2 and 3 short, last joint of
maxillary palpi acutely triangular, last joint of labial transversely lunate; 4th
joint of tarsi long lobate, claws externally divided, internally simple; three
last abdominal segments phosphorescent; last superior segment with rounded
apex.

LacorparrE, TH. Histoire naturelle des insectes, Genera des Coléop-
téres, vol. 4. Lampyres, pp. 307-340, 1857. Photuris, pp. 338-340:

Head moderately elongated or short; eyes of at least ordinary size; antennae
fairly long, most frequently very slender and bristle-like, of 11 joints, the first
a reversed cone, the 2nd and 3d of relatively variable length, the 4th often longer
than those following, these sub-equal. Prothorax transversal or not, semi-

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

circular in front, widely edged except at the.base, the angles more or less
prominent. Elytra soft, sub-parallel for the most part, more rarely oval. Legs
long and slender, posterior femora very prominent on the inner edge, Ist joint
of the posterior tarsi at least as long as the two following together, the 4th very
long, deeply divided into two slender lobes, the 5th long, in part free; claws
simple or bifid at the end. Abdomen not lobed on the sides. Body elongated,
parallel or oval, flat.

LeConrte, J. L. Trans. Amer. Ent. Soc., vol. 9, 1881. Luciolae, p. 37:

The eyes are large, convex and widely separated above and beneath in both
sexes, not conspicuously larger in ¢; the head is rounded, narrowed behind, and
not retractile; it is but partially covered by the prothorax, which is, however,
of the usual hood-like form and rounded in front. The antennae are longer
than one-half of the body, filiform, slender, not compressed, inserted near the
anterior margin of the front, and moderately approximate; the second and third
joints are about equal, and together are as long as each of the following joints.

The sexes are similar in form, with long elytra and well developed wings;
the light organs occupy the whole of the fifth and following segments; stigma-
like pores are not obvious, being situated at the base of the fifth and sixth seg-
ments and less strongly marked than in Pyractomena and Photinus g. The
seventh ventral in @ is obtusely triangular; in ¢ the fifth and sixth are broadly
emarginate, and seventh is smaller than in 9, sinuate at the sides and pro-
longed at the middle, the eighth is a little wider and longer than the prolongation
of the seventh. In our species the outer (or anterior) claw is cleft at the tip.
The prothorax and elytra are densely rugosely punctured, the former is yellow
with a black stripe or spot, each side of which the disc is red; the latter have
the whole margin and frequently a discoidal stripe pale. A single genus occurs
in our fauna with limited representation.

Oxivier, Ernst. Wytsman’s Genera insectorum, fasc. 53, p. 57,
1907:

Body elongated, parallel or oblong-oval, having a soft tegument; head hardly
visible, attenuated, on a sort of collar projecting from the prothorax; labrum
wanting or indistinct because of the proportions of the epistome; antennae long,
very slender, the second joint of variable size but always fairly long; prothorax
rounded or ogival in front, with the posterior angles sometimes obtuse and
scarcely projecting, sometimes very sharp and prolonged posteriorly; legs long
and slender; 4th joint of the tarsi bilobed, claws entire or divided... ;
abdomen composed of 7 segments, the last ones containing the luminous appa-
ratus, which is much more developed in the males. Both sexes have wings and
elytra.

... the sexual differences consist in the integrity or division of the claws,
and particularly in the ¢ the last ventral segment is short, laterally sinuate,
and terminated by a linear lobe more or less enlarged; in the ? it is large,
triangular, with an obtuse point, or slightly incised. As generally among the
Lampyridae, the eyes of the d are very large and prominent, and the head ap-
pears deeply concave.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 43

OuiviER, Ernst. Ann. Soc. Ent. France, ser. 6, vol. 6, pp. 201-240,
1886, is essentially the same as the above. He criticizes Mot-
schulsky’s splitting of Photuris into several new genera, say-
ing that it would put the two sexes of some species into separate
genera.

BRADLEY, J. C. Manual of the genera of beetles of North America,
p. 98, 1930, follows LeConte (1881) and Olivier.

RESULTS OF THE EXAMINATION OF BARBER’S SPECIMENS

2.1 brunnipennis var. floridana.

General: A small (9.5 X 3.5 mm.) lampyrid, dark brown, without dark
pronotal spot or elytral stripes.

Pronotum: Opaque white with central yellowish area; no black or
orange spots, and no sulcus. 1.8 & 2.6 mm.?

Scutellum: Yellow anteriorly, white posteriorly; rather narrower pos-
terior point than in most species; mesonotal plates yellow, rather
large.

Elytra: 7.75 1.75 mm.; rather wide lateral, and narrow sutural
margins yellow; margins continuous around tips; no stripes or
vittae.

Head: Width across eyes 1.95 mm.; eye length 0.6 mm.

Frons yellow, 0.4 mm. wide above antennal sockets, 0.75 mm. above
eyes; interocular margins rather divergent; inner edges of antennal
sockets 0.05 mm. apart.

Maxillary palpi light brown; labial palpi almost white. Labrum short,
light brown, with very narrow darker edge, and no protuberances.

Antennae: 5.1 mm. long, brown; white visible in joint sockets.

Sternites: 2 to 5 brown; 6 and 7 luminous; 8 white, with rather wide-
angled posterior point.

Legs: Coxae yellow, third pair slightly infuscate; femora yellow with
brown knees; tibiae and tarsi brown; outer claws bifid, but the
inner prong distinctly shorter than the outer one. Posterior legs
7.05 mm. long, 0.74 of the total length.

3. divisa (two specimens differing in color and slightly in size).

General: A small lampyrid (9.5-10.0 X 3.1 mm.) brown to dark brown,
with a trapezoidal median pronotal pigmentation divided longitudi-
nally by a narrow light streak partially in a narrow sulcus; elytra
with distinct white margins, but no stripe.

Pronotum: 1.75-2.0 X 2.25-2.6 mm.; central trapezoidal brown area
divided longitudinally by a narrow light line or streak, part of
which is in a narrow sulcus; in one specimen the angles are defi-
nitely produced posteriorly, in the other they are not.

Scutellum: Light brown or yellow; mesonotal plates dull brown or
yellow.

1 Numbers are those given the species in Barber’s table.
2Length and width, respectively.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 17

Elytra: 7.75-8.2 X 1.55 mm.; brown or dark brown; distinct lateral
and sutural margins white, continuous around tips; no stripes or
vittae; humeri distinctly inclined inward and backward toward
scutellum (different from other species).

Head: Width across eyes 1.85-2.0 mm.; eye length I.0 mm.

Frons brown, very wide, 0.75 mm. above antennal sockets, 0.85-0.9 mm.
above eyes; interocular margins very slightly divergent (different
from most species).

Maxillary palpi brown, labial palpi white or light brown; labrum
short, light brown.

Antennae: 4.5-5.35 mm. long, brown, unmarked, although joint sockets
may be white.

Sternites: 2 to 5 brown, 5 may be darker with narrow white posterior
edge; 6 and 7 luminous; 8 white, with a rather sharp central
point about 0.25 mm. long.

Aedeagus: 2.0 mm. long.

Legs: Coxae and femora brownish yellow, knees darker; tibiae and
tarsi brown; lobes of fourth tarsal joint relatively short. Posterior
legs 6.3-6.6 mm. long, 0.65-0.665 of total length.

4. congener (old specimen, 1914).

General: A small lampyrid (9.7 X 4.0 mm.) with parallel dark-brown
elytra without stripes, and pronotum with central yellow spot, no
black area.

Pronotum: 1.85 < 2.6 mm.; central yellow area bearing 2 indefinite
longitudinal brown streaks; sharp sulcus in anterior half.

Scutellum: Light brown; mesonotal plates yellow.

Elytra: 7.85 < 2.0 mm., brown without stripes or vittae; practically
parallel; 0.45-mm. lateral and narrow sutural margins yellow, con-
tinuous around tips.

Head: Width across eyes 2.05 mm.; eye length 1.25 mm.

Frons yellow, 0.5 mm. wide above antennal sockets, 0.95 mm. above
eyes; interocular margins more divergent than in most species;
inner edges of antennal sockets very close together, 0.05 mm.

Maxillary palpi brown, finger-tipped; labial palpi light brown, more
nearly symmetrically crescentic than in most of the species—more
like the securiform usual in Photinus.

Labrum brown, with a visible point.

Antennae: 4.95 mm. long, proportionately rather short; brown, with
white joint sockets.

Tergites: Brown, last 3 with lighter edges.

Sternites: 2 to 5 brown, posterior edge of 5 lighter; 6 and 7 luminous,
probably originally white, now yellow brown; 8 has a median
posterior point.

Legs: Coxae light brown; femora proximally light brown, shading
to dark brown at knees; tibiae and tarsi dark brown; fifth tarsal
joint appears shorter than in most species. Posterior legs 7.5 mm.
long, 0.775 of total length.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 45

5. frontalis (2 specimens which differ mainly in size).

General: A medium-sized lampyrid (12.0-13.5 X 4.2-5.2 mm.), dark,
with wide lateral elytral margins and rather short pronotum having
an indefinite brown spot.

Pronotum: 2.25-2.75 X 3.25-4.0 mm.; very short, almost semicircular ;
large central triangular ivory area, base posterior, having an in-
definite brown area; angles large and produced posteriorly about
0.25 mm. beyond median line.

Scutellum: White; mesonotal plates dull white.

Elytra: 9.0-10.6 X 2.1-2.6 mm.; brown, distinctly widened by the
0.55-mm. lateral margins, giving a somewhat oval appearance ; mar-
gins not quite continuous around tips; no stripes or vittae.

Head (larger specimen): Width across eyes 3.2 mm.; eye length
1.55 mm.

Frons ivory white, 0.7 mm. wide above antennae sockets, 1.25 mm.
above eyes; antennal sockets 0.1 mm. apart.

Maxillary palpi large, dark brown; labial palpi yellow. Labrum short,
dark brown, with 3 points or denticles.

Antennae: 7.65 mm. long in larger specimen; dark brown to prac-
tically black; joint sockets white.

Sternites: 2 to 5 brown; 6 and 7 luminous, and apparently not as much
longer than the fifth as in most species; 8 yellow, with posterior
point.

Aedeagus: 2.0 mm. long.

Legs: Coxae light brown; femora light brown proximally, darker
distally ; tibia and tarsi dark brown. Posterior legs of larger speci-
men 10.15 mm. long, 0.76 of total length.

6. pensylvanica.

General: A small lampyrid (9.0-10.0 X 3.0-3.5 mm.) with medium
brown, white-margined elytra, and pronotal black and orange pig-
mentation.

Pronotum: 1.75 X 2.25 mm.; median black or very dark brown
T-shaped area with large orange spot on each side; no sulcus.

Scutellum: Brown anteriorly, to nearly white at posterior point.

Elytra: 7.5 >< 1.6 mm.; base color brown; 0.45-mm. wide lateral and
0.3-mm. sutural margins yellow, continuous around tips; outline
nearly oval; oblique stripe from humerus about 5.0 mm. long, about
0.1 mm. wide at humerus, narrowing to end.

Head: Width across eyes 1.85 mm.; eye length 1.0 mm.

Frons ivory, 0.5 mm. wide above antennal sockets, 0.9 mm. above
eyes.

Maxillary palpi light brown; labial pale brown.

Labrum short, dark brown, with dull median point.

Antennae: 4.2 mm. long, rather short, brown; joint sockets white.

Tergites: Brown to eighth, latter ivory.

Sternites: 2 to 5 mainly light brown, posterior one-third of fifth, white;
6 and 7 luminous; 8 ivory with median point 0.25 mm. long.

Aedeagus: 1.75 mm. long.

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Legs: Coxae light and darker brown; femora mostly light or yellow-
ish brown, darker distally; tibiae and tarsi dark brown. Posterior
legs 6.8 mm. long, 0.736 of total length.

7. potomaca (two specimens, varying principally in size).

General: A small to medium-sized lampyrid (9.35-12.0 X 3.2-4.5 mm.),
light brown, subparallel, white margins and oblique stripes; black
and orange spot on pronotum.

Pronotum: 1.85-2.5 X 2.5-3.0 mm.; angles not produced posteriorly ;
median long-triangular brown mark with apex anterior, and short
triangle from this apex to anterior edge of pronotum; large orange
area on each side of brown triangle.

Scutellum: White; mesonotal plates dull dark brown.

Elytra: 7.5-9.5 X 1.6-2.25 mm.; light brown with 0.5-mm. lateral and
0.25-mm. sutural margins white; white oblique stripe 0.2 mm. wide
at humerus, narrowing to become indefinite at a length of about
5.0 mm.

Head: Width across eyes 2.25 mm.; eye length 1.2 mm.

Frons ivory white, 0.7 mm. wide above antennal sockets, 1.0 mm.
above eyes—perhaps less divergent than usual.

Maxillary palpi brown; labial light brown.

Labrum dark brown, with three dull points.

Antennae: 6.0-6.6 mm. long, brown with white rings at joint sockets ;
seventh joint somewhat the longest.

Tergites: Posterior two mainly white; others brown.

Sternites: 2, 3, and 4 brown, 5 mostly white medially; 6 and 7 lumi-
nous; 8 white with median point 0.25 mm. long.

Legs: Coxae brown; femora one-half to two-thirds yellow-brown,
distally infuscate; tibiae and tarsi darker brown; lobes of fourth
tarsal segment appear longer than usual. Posterior legs of larger
specimen 9.15 mm. long, 0.762 of total length.

8. versicolor.

General: A fairly large lampyrid (13.0-14.0 X 4.5-5.0 mm.), brown
elytra with yellow margins and yellow oblique stripe; black T and
orange pigmentation on pronotum.

Pronotum: 2.75 X 3.55 mm.; a median black or dark-brown area hay-
ing the form of a T with the cross bar lying along the posterior
edge of the pronotum; the area between the bar and foot of the
T is orange; the foot of the T connects with the slightly wider
base of a triangle, the apex of which coincides with the anterior
median line of the pronotum. Angles rounded, not produced pos-
teriorly; a row of long yellow hairs on the posterior edge of the
pronotum.

Scutellum: Brown; mesonotal plates brown.

Elytra: 11.5 X 2.5 mm., subparallel; base color brown; 0.5-mm. lateral

‘and narrow sutural margins yellow, continuous around tips; yel-
low oblique stripe 0.25 mm. wide, not appreciably wider at humerus,
7.5 mm. long.

Head: Width across eyes 2.6 mm.; eye length 0.8 mm.

Frons yellow, 0.7 mm. wide above antennal sockets, 1.25 mm. above
eyes; inner edges of antennal sockets 0.1 mm. apart.

NOS ot FIREFLIES OF THE GENUS PHOTURIS—BARBER 47

Maxillary palpi and labial palpi brown.
Labrum brown; appears truncate-triangular with nearly straight edge.

Antennae: 9.2 mm. long (rather long) ; black, each joint with lighter
base; joint 3 rather longer than 2, joints 4 to 10 longer than
first joint (exceptional).

Tergite 8 appears to overlap sternite 8.

Sternites: 2, 3, and 4 brown, becoming darker in this order; 5 brown,
posterior one-third white; 6 and 7 luminous; 8 white with poste-
rior point.

Aedeagus: About 2.5 mm. long.

Legs: Coxae dark brown; femora light or yellowish brown for
proximal two-thirds, distally darker; tibiae and tarsi of anterior
two pairs of legs dark brown, of posterior pair lighter. Posterior
legs 10.5 mm. long, 0.763 of total length.

9. versicolor var. quadrifulgens.

General: Much like versicolor but darker and narrower; short, in-
distinct elytral stripe.

Pronotum: 2.6 X 3.25 mm.; pigmentation like versicolor except that
the upright of the T widens at the base to meet the base of the
terminal triangle; shallow sulcus in posterior half of the T.

Scutellum: Dark brown with lighter posterior tip; mesonotal plates
dark and light brown.

Elytra: 11.2 1.8 mm., dark brown, subparallel; 0.4-mm. lateral and
0.13-mm. sutural margins ivory, continuous around tips; oblique
light stripe and ridge from humerus to one-half elytral length.

Head: Width across eyes 2.5 mm.; eye length 1.5 mm.

Frons nearly white, 0.75 mm. wide above antennal sockets, 1.25 mm.
above eyes.

Maxillary palpi dark brown, labial dark and light brown.

Labrum short, dark brown, front edge almost straight, except for
distinct median protuberance and an indistinct one at each side.

Mandibles large and thick.

Antennae: 8.35 mm. long, practically black; proximal ends of joints
a little lighter, and joint sockets white.

Sternites: 2, 3, and 4 brown, 5 mainly brown, posterior one-third white;
6 and 7 luminous, posterior edge of 6 nearly straight; 8 white with
posterior point.

Aedeagus: 3.0 mm. long.

Legs: Coxae brown; femora yellow-brown proximally, distal two-thirds
dark brown; tibiae and tarsi dark brown. Posterior legs 9.6 mm.
long, 0.703 of total length.

10. pyralomimus (two specimens, one somewhat lighter than the one described).

General: Much like versicolor, but darker, somewhat more oval, and
with pronounced oblique stripes on elytra.

Pronotum: 2.55 < 3.8 mm.; pigmentation like versicolor; angles much
produced posteriorly; no sulcus.

Scutellum: Brown, fading to nearly white at the posterior point;
mesonotal plates dull brown.

PT.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Elytra: 10.5 X 2.45 mm., distinctly widened by the 0.5-mm. yellow
lateral margins; sutural margins 0.35 mm.; margins continuous
around tips; outline nearly oval. Oblique stripe 0.5 mm. wide at
humerus, becoming narrower, and extending almost to ends of
elytra.

Head: Width across eyes 2.5 mm.; eye length 1.2 mm.

Frons ivory, 0.9 mm. wide above antennal sockets, 1.25 mm. above
eyes.

Maxillary and labial palpi brown, latter with a low point on the
base of the thumb; tips of maxillary palpi rather wide and flat,
square-ended rather than round as usual.

Labrum dark brown with median dull point.

Antennae: 7.8 mm. long, practically black, joint sockets white.

Tergites: 3 posterior tergites white.

Sternites: 2, 3, and 4 light brown, 5 white on posterior one-third; 6 and
7 luminous; 8 white with rather sharp posterior point.

Legs: Coxae brown; femora brown for distal one-half to two-thirds,
proximally lighter; tibiae and tarsi dark brown. Posterior legs
10.35 mm. long, 0.707 of total length.

fairchildi.

General: A medium-sized lampyrid (12.0 X 4.0 mm.), light-brown,
narrow elytral margins and oblique stripes; pronotal pigmentation
similar to versicolor, but lacks the cross bar on the T.

Pronotum: 2.15 X 2.85 mm.; median black mark and orange areas
much as in versicolor, but lacks the cross bar on the T; orange
area extends nearly to the posterior margin.

Scutellum: Light brown; mesonotal plates brown.

Elytra: 9.8 X 2.0 mm., light brown; very narrow light-colored lateral
and sutural margins; narrow oblique stripe extending to within
2.0 mm. of elytral tip.

Head: Width across eye 2.15 mm.; eye length 1.3 mm.

Frons practically white, brownish under pronotum; 0.55 mm. wide
above antennal sockets, I.0 mm. above eyes; antennal sockets
0.05 mm. apart.

Maxillary palpi brown; labial, dark and light brown.

Labrum brown, filling the mandibular circle.

Antennae: 6.1 mm. long, brown, proximal ends of joints lighter.

Sternites: 2, 3, and 4 brown, 5 about one-half white; 6 and 7 luminous;
8 white with posterior point.

Aedeagus: 2.3 mm. long.

Legs: Coxae light brown; femora yellowish brown; tibiae and tarsi
mostly dark brown. Posterior legs 8.2 mm. long, 0.686 of total
length.

11a. tremulans (description prepared from two selected from a series of very

similar specimens).

General: A medium-sized lampyrid (10.25-12.5 X 4.2-4.6 mm.) ; brown
elytra with wide margins, and a short, narrow, oblique vitta on
each; pronotal pigmentation similar to that of Ph. fairchildi, but
brown area less definite. Form slightly oval.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 49

Pronotum: 2.0-2.5 & 2.55-3.0 mm., rounded ogival, posterior edge
straight but depressed just adjacent to the angles; a row of long
yellow hairs at the median posterior edge. A median long, narrow,
brown triangle, extending as a line to the median point of the
anterior edge, separates two large orange areas, and may have a
short transverse extension along the posterior edge. A 0.I-mm.
white margin, lateral and anterior, between which and the orange
area the pronotum is dense ivory-colored. No sulcus.

Scutellum: Kite-shaped, with a rather sharp posterior apex, and
angular rather than rounded anteriorly; brown, fading to yellow
at the apex. Mesonotal plates dull darker brown.

Elytra: 8.25-10.0 X 2.1-2.3 mm.; base color medium brown; lateral
margins yellow and 0.5-0.6 mm. wide; sutural margins 0.2-0.25 mm.
wide; margins continuous around elytral tips. On each elytron a
very narrow (0.1 mm. or less) pale oblique vitta from the humerus
to 0.25 to 0.4 of the elytral length. No pronounced costae.

Head: Width across eyes 2.05-2.35 mm.; eye length 1.15-1.25 mm.

Frons ivory, 0.55-0.75 mm. wide above antennal sockets, I.0-1.1 mm.
above eyes.

Maxillary palpi rather large, brown.

Labial palpi ivory to light brown, usual mitten-shape.

Labrum short, dark brown, sinuate to give three low dull pro-
tuberances.

Mandibles large, brown.

Antennal sockets white-ringed, 0.05 mm. between inner edges.

Antennae: 6.45-7.4 mm. long, practically black; yellow rings at both
proximal and distal ends of each joint; joint sockets white; first
joint longest, second shortest, third slightly longer than second,
fourth to tenth each of the same length, eleventh slightly shorter
than tenth.

Thorax: Ventrally dark brown.

Tergites: Dark brown except eighth, which is white.

Sternites: 2 to 5 mainly brown, 5 has a narrow white posterior mar-
gin; posterior edges practically straight. 6 and 7 luminous, 6 shal-
lowly and 7 more deeply notched medially, and 1.3-1.5 times as long
as 5. 8 ivory white with median point 0.25 mm. long. 9 small, ogival,
ivory white. No foveae evident.

Legs: Coxae of the first two pairs light brown, of posterior pair dark
brown; femora mainly yellow, but brownish infuscation may ex-
tend to nearly one-half length; tibia dark brown; tarsi somewhat
lighter ; lobes of fourth tarsal joint extend three-fourths length to
claws; tibial spurs large, 0-2-2. Posterior legs long, 8.6-9.5 mm.

12. caerulucens (a second specimen slightly lighter than the one described).

General: A medium-sized lampyrid (about 12.0 4.0 mm.), dark
brown; elytra rather widely margined and with long oblique stripe ;
pronotal markings like versicolor.

Pronotum: 2.5 X 3.15 mm., marked like versicolor; no sulcus.

Scutellum: Brown with white posterior tip; mesonotal plates dull
brown.

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

Elytra: 9.25 X 2.05 mm.; base color dark brown; 0.5-mm. wide lateral
and 0.2-mm. sutural margins white and continuous around tips;
oblique white stripe distinct for 5.0 mm. from humerus, becoming
indefinite.

Head: Width across eyes 2.4 mm.; eye length 1.25 mm.

Frons ivory, 0.85 mm. wide above antennal sockets, 1.05 mm. above
eyes, rather less divergent than usual.

Maxillary palpi dark brown, labial light brown; thumb of latter
pointed and curved slightly downward.

Labrum dark brown, apparently with 3 dull points.

Antennae: 6.3 mm. long, dark brown, joint sockets white.

Tergites: Brown, except 8th which is white.

Sternites: 2, 3, and 4, dark brown, 5 white on posterior one-third;
6 and 7 luminous; 8 white with rounded posterior point rather
wide-angled.

Legs: Coxae dark brown; femora and tibiae yellow brown for proximal
half, distally dark brown; tarsi darker. Posterior legs 8.95 mm.
long, 0.761 of total length.

13. aureolucens (a second specimen darker, and somewhat smaller, 10.75 X 3.9
mm., than the one described).

General: A medium-sized lampyrid (12.8 4.0 mm.), light brown
with yellow elytral margins and oblique stripe; pronotal pigmenta-
tion as in versicolor, but cross bar on T narrower and basal tri-
angle relatively larger.

Pronotum: 2.5 X 3.0 mm.; pigmentation as in versicolor, but the cross
bar on T narrower, and the triangle at the foot of the T relatively
larger. Angles slightly produced posteriorly.

Scutellum: Brown, tip white; mesonotal plates brown.

Elytra: 10.3 X 2.0 mm., base color light brown; 0.5-mm. lateral and
0.I-mm. sutural margins yellow; oblique yellow stripe covers outer
corner of humerus, narrowing rapidly to 0.2-0.25 mm., and becom-
ing indistinct at a length of 7.5 mm.

Head: Width across eyes 2.25 mm.; eye length 1.3 mm.

Frons ivory, 0.8 mm. wide above antennal sockets, 1.1 mm. above
eyes.

Maxillary palpi dark brown, labial brown.

Labrum dark brown, sinuate rather than toothed, to show three pro-
tuberances.

Antennae: 6.45 mm. long, practically black, with bases of joints
paler, and sockets white.

Tergites: Brown except last, which is white.

Sternites: 2, 3, 4, and 5 brown, a little white on posterior edge of 5; 6
and 7 luminous; 8 white with median point 0.25 mm. long.

Legs: Coxae brown; femora light yellowish brown, infuscate toward
knees; tibiae and tarsi dark brown. Posterior legs 8.85 mm. long,
0.692 of total length.

14. lucicrescens (three specimens, all light-colored, though slightly different,
and of nearly the same size and proportions).

General: A fairly large lampyrid (12.5-13.5 X 4.9-5.2 mm.), prac-
tically parallel, with margined and striped elytra and pronotal pig-
mentation resembling versicolor.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 51

Pronotum: 2.5-2.6 X 3.5-3.6 mm.; pigmentation similar to that of
versicolor but cross bar on the T very short, and orange areas
somewhat smaller.

Scutellum: White, and proportionately rather long; mesonotal plates
yellow.

Elytra: 10.25-10.8 X 2.45-2.6 mm.; base color light grayish brown, be-
coming paler toward tips; practically parallel; 0.3-0.5-mm. lateral
and 0.1-0.15-mm. sutural margins continuous around tips but in-
distinct because of pale color of elytra; oblique stripe 7.5 mm.
long from humerus.

Head: Width across eyes 2.6 mm.; eye length 1.4 mm.

Frons white, 0.85 mm. wide above antennal sockets, 1.25 mm. above
eyes.

Maxillary palpi dark brown, labial light brown.

Labrum short, white with dark brown distal edge, and an indefinite
median protuberance.

Mandibles appear large for the other proportions.

Antennae: 8.0-8.25 mm. long, very dark brown with proximal ends
of joints white, giving a beaded appearance; tenth and eleventh
joints shorter than fourth to ninth.

Tergites: Last tergite white, the two penultimate ones medially white;
remainder brown.

Sternites: 2, 3, and 4 light brown; 5 mostly white; 6 and 7 luminous;
8 ivory with triangular median point 0.35 mm. long—not as sharp
as in most species.

Aedeagus: 2.6-2.75 mm. long.

Legs: Coxae light brown to yellow; femora, tibiae, and tarsi proximally
yellow, distally brown. Posterior legs 10.1 mm. long, 0.76 of total
length.

(Specimens of this species collected in northern Delaware in 1947-48 agree
with the above except in color, being darker throughout. )
15. hiebes (a second specimen is very similar).

General: A small lampyrid (10.5 X 2.8 mm.), light brown, practically
parallel, wide lateral margins, and pronotal pigmentation some-
what resembling versicolor.

Pronotum: 2.2 X 2.7 mm., with median brown area resembling versi-
color, but lighter and less definite; orange areas similar to versi-
color.

Scutellum: White; mesonotal plates dull white.

Elytra: 8.3 X 1.4 mm., light brown; 0.5-mm. lateral and narrow sutural
margins white; margins barely continuous around tips; a very
narrow oblique white stripe on each elytron ending at about three-
fourths of the elytral length.

Head: Width across eyes 2.15 mm.; eye length 1.125 mm.

Frons ivory white, 0.6 mm. wide above antennal sockets, 1.0 mm.
above eyes; inner edges of antennal sockets 0.1 mm. apart.

Maxillary palpi brown, appearing large for this insect; labial palpi
brown.

Labrum short, brown, sinuate.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Antennae: 7.25 mm. long, dark brown with proximal ends of joints
lighter ; tenth and eleventh joints shorter than fourth to ninth.

Tergites: 6, 7, and 8 white, almost transparent; others brown.

Sternites: 2 yellow, 3 light brown, 4 darker, 5 mostly mottled white;
6 and 7 luminous; 8 white with long posterior point.

Aedeagus: 2.25 mm. long.

Legs: Coxae yellow; femora yellow-brown, knees somewhat infuscate ;
tibiae and tarsi mostly dark brown, lighter proximally. Posterior
legs 8.5 mm. long, 0.81 of total length.

16. salinus (44 specimens, including 99, available, of which 10 do were selected
as covering the range of variation. The 9? tend to be larger and
darker, and have smaller eyes and shorter antennae).

General: A medium-sized lampyrid (9-12 X 3.2-4.1 mm.), grayish to
yellowish brown under general illumination, with fairly wide, light
lateral elytral margins, narrow sutural margins, and a narrow but
distinct oblique yellow or white vitta past the midlength of each
elytron; pronotum broadly rounded to scutate with a median brown
vitta between large orange-colored areas.

Pronotum: 1.9-2.5 X 2.5-3.0 mm.; edges transparent yellow, mottled;
scutate to broadly rounded, posterior edge sinuate, but angles not
produced beyond median; a row of long yellow hairs along the
median half of the posterior edge. The pigmentation consists of a
median brown area, hourglass-shaped, 0.2 to 0.6 mm. wide at the
constriction, extending from the posterior edge nearly to the
anterior edge, sometimes narrowing to a line completely to the
anterior edge; occasionally the brown area may widen to a short
transverse bar at the posterior edge; large orange-colored areas
on each side of the brown area.

Scutellum: Transparent yellow to brown, with lighter posterior apex;
mesonotal plates the same color as the scutellum in each specimen.

Elytra: 7.5-9.25 X 1.65-2.05 mm.; base color brown to light brown,
appearing grayish or yellowish under general illumination. Distinct
lateral margins about 0.5 mm. wide, slightly widening the elytra,
giving a slightly oval outline. A narrow white to yellow oblique
vitta from the humerus to past the midlength of each elytron.
Margins continuous around the tips of the elytra, but indistinct in
lighter specimens.

Head: Width across eyes 2.1-2.4 mm.; eye length 0.9-1.35 mm.

Frons yellow, smooth or but little hairy; medially brown toward
tops of eyes. Interocular margins constricted over antennal sockets,
and divergent toward tops of eyes, intermediate edges nearly
parallel.

Maxillary palpi brown; labial palpi light brown, of the usual mitten
shape.

Labrum dark brown—practically black; distinctly tridentate in some
specimens, obscurely so in others.

Mandibles large.

Antennae: 5.55-6.35 mm. long, brown to light brown, distal ends of
joints paler, and proximal ends with a narrow light ring, giving
a distinct jointed appearance even by general illumination. Third
joint but little longer than the second.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 53

Thorax: Ventrally light to dark brown.

Tergites: 6, 7, and 8 white; the anterior ones may be white or light
brown; eighth usually rounded, but truncate-triangular in some
specimens.

Sternites: 2 to 5 light to dark brown; 5 may have a white posterior
margin; 6 and 7 luminous, white or yellow; 8 white, with a hairy
median point; 9 white, ogival.

Aedeagus: Where extruded, of the same type as in the other species.

Legs: Femora yellow, tibia and tarsi brown; tarsal spurs large, 0-2-2.
Lobes of fourth tarsal segment fairly long. Posterior pair of legs
7.0-7.9 mm. long.

17. cinctipennis (a second specimen a little longer than the one described;
otherwise similar).

General: A rather small lampyrid (10.75 < 3.5), dark brown elytra,
margined, and with narrow and short oblique stripes; pronotal
marking somewhat like versicolor.

Pronotum: 2.25 X 2.75 mm., brown pigmentation similar to versicolor,
but upright of the T very narrow and cross bar short; distinct
sulcus, widening posteriorly to include most of the short cross bar
on the T; orange area similar to versicolor.

Scutellum: Ivory white with central brown spot; mesonotal plates dull
pale brown.

Elytra: 851.75 mm., apparently a uniform dark brown except
for 0.45-mm. wide lateral and 0.2-mm. sutural margins, which
join at the rather unusually pointed tips. A very narrow light-
brown oblique stripe extends from the humerus about half the
elytral length.

Head: Width across eyes 2.1 mm.; eye length 1.2 mm.

Frons white, 0.7 mm. wide above antennal sockets, 1.0 mm. above
eyes.

Maxillary palpi light brown; labial palpi ivory, with a low pro-
tuberance on the thumb.

Labrum dull white, edge brown, with a definite median tooth, and a
duller one on each side.

Antennae: 6.65 mm. long, mostly dark brown; joints with white
proximal ends and white rings at the sockets.

Tergites: Dark brown.

Sternites: 2 to 5 mainly brown, irregularly white in posterior one-third
to one-half; 6 and 7 luminous, 6 only very slightly notched, and
7 but little more—both less than in most species; 8 white.

Aedeagus: 2.0 mm. long.

Legs: Nearly all white, hairs brown. Posterior legs 9.1 mm. long,
0.845 of total length.

18. lineaticollis (an old specimen, 1882).

General: A large lampyrid (14.5 X 5.2 mm.), very dark, margined
elytra, with pronotal pigmentation similar to versicolor, but no
cross bar on the T.

Pronotum: 2.75 X 3.6 mm., with pigmentation similar to that of versi-
color, but no cross bar on the T along the posterior edge.

Scutellum: Yellow, nearly translucent in posterior half; mesonotal
plates yellow.

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 7

Elytra: 11.8 X 2.6 mm., appear dark brown except for rather narrow
(0.35-mm.) lateral margin and (0.25-mm.) sutural margin; mar-
gins yellow, and not continuous around the tips. There is an obscure
lighter-brown oblique stripe from the humerus.

Head: Width across eyes 2.85 mm.; eye length 1.25 mm.

Frons yellow, 1.2 mm. wide above antennal sockets, 1.5 mm. above
eyes, rather wider and less divergent than usual; inner edges of
antennal sockets 0.2 mm. apart.

Maxillary palpi dark brown, labial light brown.

Labrum dark brown; a dull median tooth or protuberance, and a
sharper one on each side.

Antennae: 7.65 mm. long, mainly almost black, proximal ends of joints
lighter ; ninth to eleventh joints shorter than fourth to eighth.

Tergites: Brown.

Sternites: 2 to 5 dark brown, 5 lighter on posterior edge; 6 and 7
luminous, yellow; 8 triangular, yellow.

Legs: Coxae of first two pair light brown, of third pair very dark
brown; femora mostly dark brown, lighter proximally; tibiae and
tarsi dark brown; lobes of fourth tarsal joint rather long. Posterior
legs 10.6 mm. long, 0.73 of total length.

In the writer’s semipopular “Common Fireflies of Delaware” he ex-
pressed the idea that the species giving three to five rapid coruscations
per flash and flashing at 5- to 10-second intervals is the one which
was sent to DeGeer by Acrelius from Wilmington and described by
the former in 1774 as (Photuris) pensylvamica. The reason for
this opinion was that this is by far the commonest type of Photuris
flash now seen in the vicinity of Wilmington, although both the sharp
and crescendo flashes of lucicrescens and some of the other types de-
scribed by Barber are also present. This is a dry-land species and has
been taken in copula by the writer in a nearby wheatfield where hun-
dreds of the insects were flying over the wheat, around the border
growth, and among the trees across an adjacent road. Barber, how-
ever, calls this species Photuris versicolor Fabricius, 1798, and re-
stricts the specific name pensylvanica to a marsh species giving a two-
component flash, the first component of which is short and sharp and
the second long, basing his opinion on the probable character of the
land surface around Wilmington about 1750. He is doubtless correct
in his conjecture as to the marshes at this locality at that time; there
is still plenty of marsh land along the Delaware River and the estuary
of Christiana Creek, though most of that along the Brandywine has
been filled in. Mr. Barber did not, so far as I remember, ever , tell
me that he had arrived at this conclusion, although he did write to
me about DeGeer’s use of the word “prairies” in connection with his
description of the locale of the specimens sent by Acrelius. From the

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 55

translation of Hesselius’ Journal (Delaware History, vol. 2, No. 1,
p. 83, 1947), flying over meadows might be interpreted as “fields,”
and the “sparkling” might be more like the three- to five-flasher than
the double coruscation of Barber’s pensylvanica. I have been unable
to find out what Swedish word in Acrelius’ letters to DeGeer was
translated as “prairie’’ by the latter. Observations in both northern
Delaware and on the opposite New Jersey shore of the Delaware
River have so far failed to reveal the presence of a species giving the
double flash of Barber’s Photuris pensylvanica DeGeer, but condi-
tions here have undoubtedly changed materially in the last 200 years,
and it is not impossible that industrial wastes have exterminated a
once-plentiful species.

Free translations of the descriptions given by DeGeer and by
Fabricius are given below:

DeGeer, Hist. Ins., vol. 4, pp. 52-53, 1774:

Lampyrid elongated, elytra of a pale yellowish-gray, and thorax black in the
middle with two red spots.

Lampyris pensylvanica oblong, elytra pale grayish brick-colored, thorax black
between the margins with two red spots.

The lampyrids of this species are found in Pennsylvania. Mr. Acrelius, who
sent me them from this country, says that they are found particularly on the
prairies during the whole summer, where they glitter and appear to the eyes of the
observers as a multitude of sparks; but they sparkle even more when they fly.
One can distinguish them easily from the other species.

In size and shape they resemble the three preceding species (of lampyrids),
but the head is larger and less hidden in the thorax, which is smaller than in
the other species; there is also a greater distance between the two large black
eyes, and the antennae, which almost equal the length of the abdomen, are
slender and a little hairy. On the thorax and elytra there are many small hairs.

The disc of the thorax is pale yellow, with a large oval black spot in the
middle, beside which there are two small round red spots near the edges; the
elytra are yellowish gray, with brown shading near the anterior ends. The
abdomen is brown below, but the last three segments are sulfur yellow. The
wings are dark brown, the antennae lighter brown, and the legs ochre yellow
with some small brown spots.

Fabricius, Suppl. Entomol. Syst., p. 125, Hafnia, 17098:

L(ampyris) black, thorax spotted, elytral margins and median vitta yellowish,
apex of abdomen very light. Habitat in North America. Dom. Hirschell.

Body large; antennae black, bases of joints yellowish. Head yellowish or
black. Thorax rotund, black spot in the middle, large red spots on both sides,
and broad yellowish margin. Elytra smooth, margined with black becoming
yellowish, with abbreviated vitta. Abdomen broad, white. Legs black, knees
yellowish.

DeGeer’s description of the pronotal pigmentation sounds more like
a Photinus than a Photuris, but his mention of the partially exposed

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Ti

head and the long antennae would seem to leave no doubt of the genus
of the species described. Fabricius’ description of the pronotal pig-
mentation is more correct for Barber’s specimens of both pensylvanica
and versicolor. DeGeer fails to mention the oblique elytral stripe or
vitta ; this is quite definite in Barber’s specimen of pensylvanica, and
rather shorter in his versicolor, agreeing with Fabricius’ description.
DeGeer’s drawing is unconvincing.

Just which is pensylvanica and which versicolor, must perhaps re-
main in some doubt for the present, since neither DeGeer nor Fabri-
cius record definitely the flashes of the species they describe. Unless
further data become available, it seems well to accept Barber’s de-
cision.

LITERATURE CITED
BLatcHLey, W. S.
1924. New Coleoptera from southern Florida with notes on other interest-
ing species. Can. Ent., vol. 56, No. 7, pp. 164-170.
DEGEER, C.
1774. Mémoires pour servir a l'histoire des insectes, vol. 4, 456 pp.
DEJEAN, PIERRE F, M. A.
1833. Catalogue de la collection des coléoptéres, 3d ed.
1837. Idem, reprint, 3d ed.
Ericuson, G. F.
1847. Conspectus insectorum coleopterorum, que in Republica Peruana ob-
serva sunt. Wiegemann’s Arch. Naturg., vol. 13, pt. 1, pp. 67-185.
Fasricius, J. C.
1798. Entomologia systematica, supplementum, 572 pp.
Patt Asa,
1927. New Lampyridae. Bull. Brooklyn Ent. Soc., ns., vol. 22, No. 4,
pp. 208-211.
GorHAM, H. S.
1880. Materials for a revision of the Lampyridae. Trans. Ent. Soc. London
for 1880, Mem. 8, pp. 1-37.
Hess, WALTER N.
1920. Notes on the biology of some common Lampyridae. Biol. Bull.,
vol. 38, No. 2, pp. 39-76.
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE.
1910. Opinions rendered by. Opinions I to 25. Smithsonian Inst. Publ.
1938, pp. I-61.
1914. Ibid. Opinions 57 to 65. Smithsonian Inst. Publ. 2256, pp. 131-160.
INTERNATIONAL RULES OF ZOOLOGICAL NOMENCLATURE.
1926. In Proc. Biol. Soc. Washington, vol. 39, pp. 75-103.
LACORDAIRE, J. TH.
1857. Histoire naturelle des insectes. Genera des Coléoptéres, vol. 4.
Laporte, F. L. bE.
1833. Essai d’une révision du genera Lampyre. Ann. Soc. Ent. France,
vol, 2, pp. 122-153.

NO. I FIREFLIES OF THE GENUS PHOTURIS—BARBER 57

LeConrte, J. L.
1849. Coleopterous insects. Jn White’s Statistics of Georgia, supplement
(Catalogue of the fauna and flora of the State of Georgia), pp.
25-36.
1850. General remarks upon the Coleoptera of Lake Superior. Jn Agassiz,
Lake Superior, p. 228.
1852. Synopsis of the lampyrides of temperate North America. Proc.
Acad. Nat. Sci. Philadelphia, vol. 5, pp. 331-347. (Usually incor-
rectly dated 1851.)
1881. Synopsis of the Lampyridae of the United States. Trans. Amer. Ent.
Soc., vol. 9, pp. 15-272.
Lena, C. W.
1920. Catalogue of the Coleoptera of America, north of Mexico. x + 470 pp.
Lenc, CHARLES W., and MutTcHLeEr, ANDREW J.
1922. The Lycidae, Lampyridae and Cantharidae (Telephoridae) of the
West Indies. Bull. Amer. Mus. Nat. Hist., vol. 46, art. 8, pp. 413-
499.
LINNAEUS, C.
1767. Systema naturae, 12th ed., vol. 1, pt. 2, pp. 533-1327.
McDermott, F. A.
1917. Observations on the light-emission of American Lampyridae. Can.
Ent., vol. 49, No. 2, pp. 53-61.
MELSHEIMER, F. E.
1845-1846. Descriptions of new species of Coleoptera of the United States.
Proc. Acad. Nat. Sci. Philadelphia, vol. 2, No. 12, pp. 302-318.
MorscHUuLsky, V.
1853. Etudes entomologiques, Ann. 1, 1852.
1854. Idem, Ann. 2 and Ann. 3.
1855. Idem, Ann. 4.
MutcHier, ANDREW J.
1923. Notes on West Indian Lampyridae and Cantharidae (Coleoptera)
with descriptions of new forms. Amer. Mus. Nov., No. 63, pp. 1-9.
OtivieErR, E.
1886. Etudes sur les Lampyrides. Ann. Soc. Ent. France, ser. 6, vol. 6,
Ppp. 201-246.
1899. Les Lampyrides typiques du muséum. Bull. Mus. Hist. Nat., Paris,
vol. 5, pp. 371-373.
1907. Family Lampyridae. Wytsman’s Genera insectorum, fasc. 53, Coleop-

tera, 74 pp.
1910. Lampyridae. Junk-Schenkling, Coleopterorum catalogus, pt. 9, pp.
1-68.
Outvirr, G. A.
1790. Entomologie, ou Histoire naturelle des insectes . . . Coléoptéres,

vol. 2, pt. 28.
Panzer, G. W. F.
1789. Einige seltene Insecten. Naturforscher, vol. 24, pp. 1-35.
SHarp, D., and Mur, F.
1912. The comparative anatomy of the male genital tube in Coleoptera.
Trans. Ent. Soc. London, 1912, pp. 477-642, pls. 42-78.

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Sorter, A. J. J.
1849. In Gay, Historia fisica y politica de Chile, vol. 4, p. 445.
Sturm, J.
1843. Catalog der Kafer-Sammlung, xii + 386 pp.
WENZEL, H. W.
1896. Notes on Lampyridae, with the description of a female and larva.
Ent. News, vol. 7, No. 10, pp. 294-206.
WitiaMs, F. X.
1917. Notes on the life-history of some North American Lampyridae.
Journ. New York Ent. Soc., vol. 25, No. 1, pp. 11-33.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 2

ADDITIONAL FORMS OF BIRDS FROM
COLOMBIA AND PANAMA

BY
ALEXANDER WETMORE

Secretary, Smithsonian Institution

2200000092

(Pustication 4052)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 25, 1951

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ADDITIONAL FORMS OF BIRDS FROM
COLOMBIA AND PANAMA

By ALEXANDER WETMORE
Secretary, Smithsonian Institution

As work on our ornithological collections from Colombia and Pan-
ama has proceeded, several additional forms new to science have been
found and are described in the following pages. There are included
with these a few other reports for kinds of birds only partly or not
previously known from the two republics concerned, to place this
information on record for the benefit of others.

Family ARDEIDAE: Herons
BUBULCUS IBIS IBIS (Linnaeus)
Ardea Ibis LINNAEUS, Syst. Nat., ed. 10, vol. 1, 1758, p. 144 (Egypt).

A female cattle egret was taken by M. A. Carriker, Jr., at Punto
Muchimbo, Valle, on the lower Rio San Juan about 3 kilometers be-
low Palestina, and on the opposite side of the river, on January 3,
1951. The bird is in immature dress, with only a small amount of
buff on the crown. This apparently is the first record for Colombia
and is indication that this Old World species is now spreading widely
over northern South America. It is now recorded from Dutch Gui-
ana, British Guiana, Venezuela, and Colombia. Observers should
watch for small white herons, in general like the snowy heron, but
with yellow bill. In adults the bill is described as reddish at the base,
but these would stand out otherwise because of the strong buff mark-
ings on crown and breast.

Family COCHLEARIIDAE: Boart-BiLLep Herons
COCHLEARIUS COCHLEARIUS COCHLEARIUS (Linnaeus)

Cancroma cochlearius LINNAEUS, Syst. Nat., ed. 12, vol. 1, 1766, p. 233
(Cayenne).

A boat-billed heron that I shot at the mouth of Rio Imamado on
the upper Rio Jaqué, Darién, on April 16, 1947, is a specimen of the
pale-colored, typical form, not reported previously in Panama.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 2

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

COCHLEARIUS COCHLEARIUS PANAMENSIS Griscom

Cochlearius zeledoni panamensis Griscom, Amer. Mus. Nov., No. 235, November
II, 1926, p. 11 (Corozal, Canal Zone).

A female collected by M. A. Carriker, Jr., at Acandi, Choco, on the
western shore of the Golfo de Uraba, January 17, 1950, is representa-
tive of this dark-plumaged form of Panama, here recorded for the
first time from Colombia. It will be interesting to determine if this
subspecies extends into the lower Atrato Basin, particularly in view of
the record of the typical race given above in extreme southwestern
Darién.

Family CAPRIMULGIDAE: GoatsuckKErRs
CHORDEILES ACUTIPENNIS MICROMERIS Oberholser

Chordeiles acutipennis micromeris Oberholser, U. S. Nat. Mus. Bull. 86, April 6,
1914, pp. 24 (in key), 100 (Xbac, Yucatan).

In a series of nighthawks collected January 17, 1947, at El Dificil,
Magdalena, in the level country to the east of the lower Rio Mag-
dalena I find six specimens of this form, previously unrecorded in
Colombia. Another was taken at Camp Costa Rica, not far distant,
January 26. Others were shot at Norosi, March 14, 1947, and at La
Raya, January 22, 1948, both localities being in Bolivar. These birds
are wholly typical of this form of Central America, which is common
in migration in Panama and apparently also in northern Colombia.

Family APODIDAE: Swirts
CHAETURA SPINICAUDA AETHERODROMA, new subspecies

Characters——Similar to Chaetura spinicauda fumosa? but smaller.

Description—U.S.N.M. No. 409438, male, Chepo, from 500 feet
elevation on Cerro Carbunco, Provincia de Panama, Panama, col-
lected April 14, 1949, by A. Wetmore and W. M. Perrygo (orig. No.
14808). Crown, back, wings, tail, and upper tail coverts dull black,
with a very faint greenish cast; rump pale smoke gray, shading to
smoke gray on lower back; sides of head and of neck hair brown; an
indistinct spot of smoke gray in front of eye ; throat somewhat whiter
than pale smoke gray; upper breast mouse gray, shading to deep
mouse gray on abdomen and to dark mouse gray on sides, flanks, and

1 Chaetura fumosa Salvin, Proc. Zool. Soc. London, 1870, p. 204 (Bugaba,
Chiriqui).

NO.2 FORMS OF BIRDS FROM COLOMBIA AND PANAMA—-WETMORE 3

under tail coverts; under wing coverts dark mouse gray. Bill black;
tarsus chaetura black ; claws chaetura drab (from dried skin).

Measurements.—Males, 13 specimens, wing 100.2-105.2 (103.0),
tail 36.8-40.5 (38.7), culmen from base 4.8-5.8 (5.2), tarsus 10.0-
11.6 (10.6 ) mm.

Females, 4 specimens, wing 100.3-107.8 (104.0), tail 37.7-42.7
(40.6), culmen from base 5.0-5.7 (5.2), tarsus II.0-11.5 (11.1) mm.

Type—Male, wing 101.3, tail 40.0, culmen from base 5.8, tarsus
10.0 mm.

Range.—Panama, from the Provincia de Panama (Chepo, Charco
del Toro on Rio Majé) through the Comarca de San Blas (Permé)
and Darién (Cana, Jaqué) to Colombia, in Antioquia (Puerto Val-
divia, Novita, El Real), Cauca (Juntas de Tamana, Rio San Juan),
and northeastern Magdalena (Cacagualito).

Remarks.—Small swifts of the species Chaetura spinicauda are
common in many localities but are so difficult to obtain that it has
taken several seasons in the field to secure a sufficient series to de-
termine the identity of those found in Panama and northern Colom-
bia. When enough had been collected it became apparent that two
groups differing in size were included in the range currently assigned
to the race fumosa, a larger one in Costa Rica and Chiriqui, and a
smaller one in eastern Panama and Colombia. As the form was de-
scribed by Salvin from two specimens from Bugaba, western Chiriqui,
it has been necessary first to ascertain the measurements of this
original lot. This has been possible through the kind cooperation of
J. D. Macdonald of the British Museum (Natural History), who has
verified the information that the two skins had been collected by Arcé,
and says further that a definite type had not been selected between
them. He has furnished measurements of both right and left wings,
for the chord, and with the wing flattened. One specimen had the tip
of the left wing much worn. The chord of the right wing in these
two is 109.0 and 108.0 mm., measurements that fall within the limits
of the larger group, as is shown by the following data from Costa
Rican skins. The series includes those in the Museum of Comparative
Zoology and in the Carnegie Museum, which I have examined through
the assistance of J. L. Peters and W. E. Clyde Todd.

Males, 8 specimens, wing 108.5-115.4 (110.7), tail 37.7-40.7 (38.9),
culmen from base 4.8-5.5 (5.1), tarsus 9.9-11.8 (10.6) mm.

Females, 8 specimens, wing 105.5-113.4 (109.9), tail 39.3-41.5
(39.8), culmen from base 4.6-6.0 (5.3), tarsus 9.6-11.6 (10.6) mm.

Specimens of the typical form examined from Costa Rica come

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

from Pozo Azul de Pirris and El General. Intergradation between
the two races is assumed to take place between eastern Chiriqui and
the western part of the Province of Panama, an area from which no
specimens are at present available to me.

Hellmayr,” in discussing fumosa, considered the type to be “Arcé
coll. Nr. 3328,” which is the bird noted above as having the left
wing much abraded at the tip. It is evidently this worn left wing that
Hellmayr measured to arrive at the length of 106 mm. that he lists
in the reference cited, since Mr. Macdonald gives 107 mm. for fiat-
tened left wing in this same bird. The right wing, on the contrary,
Macdonald writes, measures 112.5 mm. flattened, which is in line with
Hellmayr’s measurements for the other specimen from the type lo-
cality.

Family TROCHILIDAE: Hum™Mincsirps
HYLOCHARIS GRAYI HUMBOLDTII (Bourcier and Mulsant)

Trochilus humboldtii BourcierR and Mutsant, Ann. Sci. Phys. Nat. Agr. Ind.,
Soc. Roy. Agr. Lyon, sér. 2, vol. 4, 1852, p. 142 (Rio Mira, Esmeraldas,
Ecuador).

A male and two females that W. M. Perrygo and I collected near
Jaqué on the coast of Darién on March 18 and 26 and April 11, 1946,
are the first reported north of the Baudo region of northwestern
Colombia.

Family FURNARIIDAE: Ovenstirps
XENERPESTES MINLOSI UMBRATICUS, new subspecies

Characters —Similar to X enerpestes minlosi minlosi Berlepsch * but
decidedly darker above, without definite light streaks on the crown;
hindneck, back, wings, and tail darker.

Description —Type, U.S.N.M. No. 443152, male, Rio San Juan at
Punto Muchimbo, Valle, Colombia, December 24, 1950, M. A. Car-
riker, Jr. (orig. No. 19079). Anterior half of crown dull black,
shading gradually to blackish mouse gray across the posterior section
of the pileum to the hindneck, the feathers, except on the forecrown,
edged slightly with deep mouse gray; back and scapulars dark olive-
gray; rump and upper tail coverts light grayish olive; primaries and
secondaries sooty black, edged lightly with hair brown; wing coverts

2Verh. Orn. Ges. Bayern, 1907, vol. 8, 1908, p. 161.
3 Xenerpestes minlosi Berlepsch, Ibis, 1886, p. 53, pl. 4 (near Bucaramanga,
Colombia).

NO. 2 FORMS OF BIRDS FROM COLOMBIA AND PANAMA—WETMORE 5

blackish mouse gray, the middle and greater series tipped with white
spots that form two well-marked wing bars; middle rectrices and
outer webs of others deep olive-gray, the concealed portions of the
outer ones deep mouse gray, the two outermost edged lightly at the
tip, mainly on the inner web, with pale olive-gray; lores olive-buff,
beginning at the nostril, becoming dull white, and passing backward
above the eye as a prominent superciliary stripe; sides of head deep
olive-buff anteriorly, passing into mouse gray posteriorly ; under sur-
face dull whitish, washed indistinctly with primrose yellow, with a
few small, indistinct flecks of dark neutral gray on the sides of the
upper breast; bend of wing and axillars marguerite yellow; under
wing coverts white; inner webs of primaries edged with dull white;
under tail coverts olive-buff. Maxilla dark neutral gray ; tip of mandi-
ble neutral gray, base grayish olive ; tarsus and toes deep neutral gray,
claws whitish. (From dried skin.)

Measurements——Males, 2 specimens, wing 57.1-57.3 (57.2), tail
41.6-45.6 (43.6), culmen from base 13.4 (one specimen only), tarsus
15.3-16.0 (15.6) mm.

Females, 2 specimens, wing 52.5-54.9 (53.7), tail 42.2-42.6 (42.4),
culmen from base 12.8-13.7 (13.2), tarsus 14.8-15.1 (14.9) mm.

Type.—Male, wing 57.1, tail 41.6, (extreme tip of culmen broken),
tarsus 16.0 mm.

Range.—From eastern Darién, Panama (Garachiné, Rio Sambu,
Cituro), to northern Valle, northwestern Colombia (Malaguita, Punto
Muchimbo). (A record from Tierra Alta on the lower Rio Sint,
western Bolivar, may refer to this form. )

Remarks.—The four specimens of the new race seen have been
compared with two topotypes of minlosi in the U. S. National Mu-
seum from Hacienda Santana, above Bucaramanga, Santander. The
two minlosi are definitely paler above, and have the fore part of the
crown clearly streaked with whitish, a character noted in the original
description and shown in the plate accompanying it. It is supposed
that the records from Boyaca and Cundinamarca pertain to the typical
subspecies, while that from Tierra Alta in western Bolivar may be
the new form.

The type specimen of wmbraticus was shot high up in a tall tree,
on the south side of the Rio San Juan, near the mouth of Rio Calima.

As further specimens have come to hand from widely scattered
localities it has begun to seem probable that this bird is more common
than has been supposed. Because of small size and lack of striking
markings it is often overlooked.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS, LZ,

PREMNOPLEX BRUNNESCENS MNIONOPHILUS, new subspecies

Characters.—Similar to Premnoplex brunnescens distinctus Gris-
com * but definitely grayer ; back, rump, and upper tail coverts more
olive, less rufescent; crown slightly grayer; under surface lighter,
grayer, with throat and other markings lighter buff.

Description.—Type, U.S.N.M. No. 443674, male, south face of
Cerro Campana, Provincia de Panama, Panama, at 3,000 feet eleva-
tion, March 19, 1951, A. Wetmore and W. M. Perrygo (orig. No.
16437). Crown chaetura black, each feather being lighter centrally,
the light markings being dull colonial buff on the forehead, becoming
progressively duller posteriorly, shading to olive-buff in center of
crown, and to citrine drab on nape, the feathers therefore with a squa-
mate appearance; lores chamois, mixed with bristly projecting fila-
ments chaetura black in color; a somewhat indistinct superciliary
chamois; auricular region chaetura drab, with indistinct shaft streaks
of chamois; feathers on hindneck very dull colonial buff centrally,
forming an indistinct collar; back dull snuff brown, with faintly indi-
cated darker distal edgings, producing indistinct squamations ; rump
and upper tail coverts Prout’s brown; lesser wing -coverts sepia;
middle and greater wing coverts Prout’s brown; primaries and secon-
daries chaetura black, edged broadly on outer web with Prout’s brown ;
rectrices bister with fuscous shafts ; throat chamois ; chin paler, nearer
cream-buff, the feathers tipped very faintly with dusky neutral gray ;
feathers of upper breast cream-buff, with well-marked edgings of
dark olive, producing prominent light spots; lower breast and abdo-
men deep olive, with elongate, tear-shaped central spots of cream-buff,
these becoming progressively narrower and less in size until they dis-
appear on the lower abdomen; sides and flanks sepia, the sides with
a few elongate central markings of cream-buff; under tail coverts
Prout’s brown, with indistinct spots of buckthorn brown; under wing
coverts deep olive, mixed with chamois ; edge of wing, at base of outer
primary, chamois barred with deep olive. Maxilla blackish ; mandible
pale olive-buff with a blackish line along tomium ; tarsus and toes fus-
cous ; claws light drab (from dried skin).

Measurements—Males, 2 specimens, wing 61.2-61.6 (61.4), tail
55-9-50.2 (56.1), culmen from base 14.6-15.6 (15.1), tarsus 18.9-19.1
(19.0) mm.

Female, 2 specimens, wing 60.0-60.4 (60.2), tail 56.1-56.8 (56.4),
culmen from base 16.0-16.3 (16.1), tarsus 19.1-19.6 (19.3) mm.

4 Premnoplex brunnescens distinctus Griscom, Amer. Mus. Nov., No. 280,
September 10, 1927, p. 5 (Chitra, 4,000 feet elevation, Veraguas, Panama).

NO. 2 FORMS OF BIRDS FROM COLOMBIA AND PANAMA—WETMORE of

Type.—Male, wing 61.2, tail 55.9, culmen from base 15.6, tarsus
19.1 mm.

Range.—Known only from 3,000 feet elevation on the more humid
areas of Cerro Campana, Provincia de Panama, Panama.

Remarks.—The new form here described differs from Premnoplex
brunnescens albescens Griscom, of the mountains of eastern Darién,
in being more olive, less rufescent above, and definitely more buffy on
the throat and other light markings of the lower surface. In general
the new race is more or less intermediate in appearance between its
two geographically nearest relatives, distinctus of Veraguas and
albescens of Darién, but differs from both in being more olive, less
rufescent above. All three are closely similar in size.

SCLERURUS GUATEMALENSIS ENNOSIPHYLLUS,® new subspecies

Characters —Similar to Sclerurus guatemalensis salvini Salvadori
and Festa,® but lighter, grayer above; paler below; outer webs of
primaries more olive.

Description—Type, U.S.N.M. No. 392775, female, from Volador
(near El Tigre), 2,600 feet elevation, 25 miles west of Simiti, Bolivar,
Colombia, May 27, 1947, by M. A. Carriker, Jr. (orig. No. 11123).
Crown, hindneck, and upper back sepia, the forehead tipped lightly
with isabella color; lesser and middle wing coverts and lower back
bister ; lower rump and upper tail coverts chestnut-brown; outer webs
of greater wing coverts, primaries and secondaries Mars brown; inner
webs fuscous; rectrices sooty black; throat white, with the feathers
edged with dusky neutral gray, producing a scalloped appearance ;
sides of head sepia, with slight shaft streakings of isabella color, which
become broader in a line from below rictus back under the auricular
region, producing a faint streak ; sides of neck mingled sepia and Sac-
cardo’s umber; feathers of lower foreneck and upper breast sayal
brown centrally, tipped with bister, with shaft lines of cinnamon-buff,
producing an indistinct spotting ; lower breast and sides between Sac-
cardo’s umber and sepia; abdomen and under tail coverts between
bister and sepia; under wing coverts tawny-olive ; inner webs of cen-
tral primaries edged with avellaneous on under surface. Maxilla
brownish black; tip of maxilla fuscous, base buffy brown; tarsus

5 From évvocigvAdos, shaking or scattering leaves, in allusion to the active
habit of this bird in overturning and throwing dead leaves about in search for
food.

8 Sclerurus salvim Salvadori and Festa, Boll. Mus. Zool. Anat. Comp. Torino,
vol. 14, No. 362, November 17, 1899, p. 23 (Rio Peripa, Ecuador).

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

blackish brown, becoming buffy brown on upper end of posterior face ;
feet blackish brown, claws slightly browner (from dried skin).

Measurements.—Males, 9 specimens, wing 85.3-90.1 (87.5), tail
38.2-64.2 (60.9), culmen from base 22.6-24.9 (23.8), tarsus 21.4-
23.9 (22.1) mm.

Females, 5 specimens, wing 85.8-90.0 (87.9), tail 56.8-59.0 (57.6),
culmen from base 22.0-23.9 (22.7), tarsus 21.4-22.6 (22.1) mm.

Type—Female, wing 88.3, tail 57.6, culmen from base 22.6, tarsus
22.5 mm.

Range-—From the Rio Sint valley (Nazaret, Tierra Alta, Que-
brada Salvajin) eastward through the foothill area to the western
slopes above the lower Rio Magdalena (Santa Rosa, Volador), Boli-
var, Colombia.

Remarks.—Description of the present form marks a considerable
extension of range for this species through north-central Colombia,
as previously these birds have been reported only from the west-coast
area and the upper reaches of the Rio Sint. The three geographic
races now recognized are based on small differences in color that
change almost insensibly over intermediate areas of considerable ex-
tent. No appreciable variation in size is evident in the three forms.
The type race, which is brighter, more reddish brown above, extends
from Guatemala (where few have been found to date) to Panama,
being found in typical form to the Canal Zone. Beyond this area there
is a gradual change through eastern Panama Province and Darién to
the darker, more sooty salvini, which ranges on the Pacific slope from
eastern Darién to Ecuador. The previously unrecognized form de-
scribed here on basis of more grayish olive color, extends eastward
in Colombia from the Sint valley through the forested hill country
of southern Bolivar to the western side of the Magdalena valley west
of Simiti. This race intergrades with salvini in the eastern part of
the Rio Atrato valley, specimens from Villa Artiaga, near Pavaron-
docito, being intermediate. The record from Quimari on the upper
Sint, reported by de Schauensee* under the name Sclerurus guatema-
lensis guatemalensis, belongs without question under ennosiphyllus, as
the typical race extends to the south and east only to western Darién.

SCLERURUS MEXICANUS OBSCURIOR Hartert

Sclerurus mexicanus obscurior Hartert, Nov. Zool., vol. 8, October 5, 1901, p. 370
(Lita, Esmeraldas, Ecuador).

Three collected April 17, 19, and 24, 1912, by E. A. Goldman at the
head of Rio Limon, on Cerro Pirri, Darién, agree fully with birds

7 Caldasia, vol. 5, No. 24, July 10, 1950, p. 690.

NO. 2 FORMS OF BIRDS FROM COLOMBIA AND PANAMA—WETMORE 9

from extreme western Colombia and northwestern Ecuador. This
race has not been reported previously north of the Province of Valle,
western Colombia.

Family FORMICARIIDAE: Awnt-THRUSHES
MYRMORNIS STICTOPTERA (Salvin)

Rhopoterpe stictoptera SAtvin, Bull. Brit. Orn. Club, vol. 1, No. 6, March 1,
1893, p. 32 (Santo Domingo, Nicaragua).

Peters ® gives the first record of this bird for Colombia from El
Umbo, Boyaca, the specimen being in the Museum of Comparative
Zoology. M. A. Carriker, Jr., collected a pair at Socarré, Bolivar, on
the Rio Sint, April 21 and 22, 1949, and another pair at Unguia,
Choco, March 10 and 16, 1950. On careful comparison with an ex-
cellent series from Nicaragua I find no differences in these birds
from points far distant from the type locality.

All material that I have seen is so clearly distinct from VM. torquata
that I consider stictoptera and torquata specifically separated.

Family FRINGILLIDAE: Grosseaks, FINCHES, BUNTINGS
ZONOTRICHIA CAPENSIS ORESTERA, new subspecies

Characters.—Similar to Zonotrichia capensis costaricensis Allen,°
but decidedly darker, less buffy brown above; dark markings of dorsal
surface blacker; gray of crown darker; chestnut of nuchal collar
deeper ; paler edgings darker; rump darker gray; tail darker; sides
and flanks grayer, less buffy ; under surface, in general, whiter.

Description—Type, U.S.N.M. No. 434103, male adult, southeast
face of Cerro Campana, Provincia de Panama, Panama, 2,000 feet
elevation, March 2, 1951, A. Wetmore and W. M. Perrygo (orig.
No. 16122). Median crown stripe and auricular area, extending for-
ward toward gape, mouse gray; superciliary stripe light mouse gray,
changing to pale mouse gray behind the eye; a few whitish feathers
behind nostril; broad lateral crown stripes, an indistinct circle around
eye, and indistinct edgings on gray auriculars and rictus dull black;
circlet of feathers on eyelids dull whitish with whitish bases on
feathers immediately in front of eye; hindneck and sides of neck
Mikado brown, forming a distinct collar ; feathers of back and scapu-

8 Check-list Birds of the World, vol. 7, 1951, p. 256.
9 Zonotrichia capensis costaricensis Allen, Bull. Amer. Mus. Nat. Hist., vol. 3,
September 29, 1801, p. 374 (San José, Costa Rica.)

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

lars black centrally, forming heavy longitudinal streaks, bordered
broadly with snuff brown laterally and tipped with deep olive-buff ;
rump hair brown; upper tail coverts olive-brown; lesser wing coverts
sooty black, edged lightly with mouse gray; middle coverts similar
but tipped prominently on the distal part of the outer web with white,
forming a distinct bar; greater wing coverts olivaceous-black, edged
with snuff brown, and tipped lightly with white to form a second bar ;
primaries and secondaries fuscous-black, edged lightly with pale smoke
gray; tertials dull black, margined with snuff brown; rectrices fus-
cous-black ; throat pure white; sides of foreneck black, the two sides
united across the front by an indistinct band where the feathers are
white tipped with deep to dusky neutral gray; breast paler than pale
olive-gray ; abdomen white; under tail coverts pale olive-buff; sides
drab, becoming slightly more buffy brown on flanks; under wing
coverts light grayish olive, edged with dull white; edge of wing dark
mouse gray, with a distinct spot of white at base of ninth primary ;
inner webs of primaries edged on lower surface with a line of pale
olive-buff. Bill dull black, with a wash of fuscous at base of man-
dible ; tarsus and toes buffy brown; claws fuscous (from dried skin).

Measurements.—Males, 8 specimens, wing 60.5-64.6 (62.9), tail
50.4-54.9 (52.6, average of 7 individuals), culmen from base 12.2-
13.2 (12.8, average of 7 individuals), tarsus 21.0-22.7 (21.6) mm.

Female, 1 specimen, wing 59.5, tail 51.6, culmen from base 12.6,
tarsus 21.2 mm.

Type.—Male, wing 63.2, tail 52.1, culmen from base 13.2, tarsus
21.0 mm.

Range.—Open slopes of Cerro Campana, in the western part of
Provincia de Panama, Panama.

Remarks.—lIt is only recently that these birds have been noted in
the mountains immediately west of the Canal Zone, and the series
recorded here includes the first specimens that have been taken in
this section, so far as I am aware. The type locality of orestera in an
airline is distant only 50 kilometers from Balboa.

The birds range over open, grass-covered ridges where exposures
of volcanic rock thrust out from the poor, stony soil, and woody
vegetation is composed of low, scrubby growth in sheltered valleys,
enly scattered shrubs appearing elsewhere. The sparrows remain
mainly in the grassland, perching on boulders or in the low bushes.
They were found from 1,800 feet upward. Their haunt is now easily
accessible, as it is possible to drive in a passenger car to the slopes
where the birds are common. In fact this was one of the first birds

NO. 2 FORMS OF BIRDS FROM COLOMBIA AND PANAMA—WETMORE II

that I saw on my visit to this mountain. While inconspicuous, these
sparrows are not especially shy, so that the species is one that is
easily found when its proper haunts are known.

The extent of the range is at present uncertain. Three specimens
in the American Museum of Natural History from Calobre and
Santa Fé in eastern Veraguas are to be placed with orestera, though
slightly intermediate toward costaricensis. An old skin in the U. S.
National Museum marked Cascajal, Coclé, appears nearer costari-
censis. Presumably this was taken on the high open slopes on the
Pacific side, above the head of the Rio Cascajal, which is a tributary
of the Rio Coclé del Norte. Dr. Matthew W. Stirling, who has been on
the Cascajal, informs me that the Caribbean slope is heavily forested,
but that there are open, grassy slopes just over the divide toward La
Pintada. All specimens from Chiriqui that I have seen are to be
placed definitely with costaricensis, so that the area of intergradation
between the two forms would appear to be in Veraguas. A sight
record from Cerro Chame by Gerald Rogers *° presumably refers to
orestera, since this mountain lies adjacent to Cerro Campana a few
miles to the southeast, the two being separated by a low divide.

10 Auk, 1945, p. 641.

| Neal
i ae > LAN

a ne i ay
hihi y's Ie

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 3

RELATIONSHIPS OF CERTAIN GENERA
Co PUNGUS GNATS. OF THE
PoMwMliy sMYCE POPHILIDAE

BY
F. R. SHAW anv M. M. SHAW

Ambherst, Mass.

(Pustication 4053)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 27, 1951

The Lord Baltimore Press

BALTIMORE, MD., U. 8 A.

RELATIONSHIPS OF CERTAIN GENERA OF
FUNGUS GNATS OF THE FPANMEY
MYCETOPHILIDAE

By F. R. SHAW anp M. M. SHAW1
Amherst, Mass.

The present study represents a continuation of a preliminary in-
vestigation of the possible value of thoracic sclerites in determining
the relationships of certain insects. Dr. G. C. Crampton was the
first to demonstrate the use of these sclerites as a means of determin-
ing the systematic position of insects. In 1925, he published a clas-
sical study of the comparative morphology of the thorax of nontipu-
loid Nematocera. In 1948 Shaw presented a paper in which he indi-
cated the value of thoracic sclerites as an aid in determining the phy-
logeny of the Mycetophilidae. Although the number of genera he
studied was admittedly small, principles were developed that have
been of value in distinguishing the phylogenetic relationships of cer-
tain genera.

Edwards (1925) was the first to indicate that the structure of
thoracic sclerites might be of value in determining generic characters
in this group. In his monograph of the British fungus gnats he noted
that in certain genera the sclerites differed in form and that such dif-
ferences might be of value in separating groups of these insects.

1 We wish to express our thanks to the Society of Sigma Xi for a grant-in-
aid that made possible the preparation of the illustrations for this paper. Also,
in the progress of this research, several others have been of invaluable assistance.
To Elmer Smith much credit is due for preparing the figures and for his keen
interest and observations. To Dr. John Lane, of Sao Paulo, Brazil, we are in-
debted for specimens of certain genera and for helpful suggestions. Dr. E. G.
Fisher offered pertinent suggestions relating to the phylogeny of the group.
Dr. Paul Freeman, of the British Museum of Natural History, kindly lent cer-
tain specimens for examination.

Owing to the comparative scarcity of specimen material of some of the fungus
gnats of this family, it was not possible in all cases to prepare specimens prop-
erly for morphological study. Consequently, no figures were made for the
genera Manota, Lygistorhina, Stenophragma, Platyroptilon, and Allactoneura,
but sufficient observations were made from pinned specimens in most instances
to determine the generic affinities.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 2

z SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

In Shaw’s 1948 paper 21 mycetophilid genera were figured and
discussed. He indicated the features that appeared primitive and
using these as principles was able to indicate the relationships of the
forms studied from a phylogenic standpoint.

Of the genera studied, 45 are figured herein. While thoracic
sclerites are primarily used, in some cases venation and chaetotaxy
are also considered. Representatives of all but two subfamilies, the
Lygistorhininae and the Manotinae, are figured. On the basis of the
present investigation, certain genera are no longer considered as dis-
tinct. In some instances the position of certain genera within tribes
is questioned, and two new tribes are proposed. Certain genera for-
merly united to others are recognized as distinct on the basis of
thoracic sclerites.

To aid the reader in visualizing the scope of the work the following

table is presented. The position of the genera is indicated as they are’

placed as a result of these studies. Genera indicated by an asterisk
are not figured but have been studied. A question mark indicates that
the affinities of the genus are still somewhat uncertain.

Subfamily Tribe Genera represented
Bolitophilinae Bolitophila Meigen (in-
cluding Bolitophilella
Landrock)
Ditomyiinae Centrocnemis Phillipi ?

Nervijuncta Marshall
Symmerus Walker
Calliceratomyia Lane

Diadocidinae Diadocidia Ruthe

Ceroplatinae (including Palaeoplatyura Meunier
Macrocerinae: genera Proceroplatus Edwards
Macrocera and Fen- Ceroplatus Bosc
deromyia) Platyroptilon Westwood *

Apemon Johannsen ?
Platyura Meigen
Macrocera Meigen
Fenderomyia Shaw
Sciophilinae Mycomyiini Mycomyia Rondani
Sciophilini Eudicrana Loew
Monoclona Mik *
Neuratelia Rondani
Parvicellula Marshall
Phthinia Winnertz
Polylepta Winnertz
Sciophila Winnertz
Stenophragma Skuse *
Syntemna Winnertz

NO. 2 FUNGUS GNATS—SHAW AND SHAW 5

Subfamily Tribe Genera represented

Sciophilinae Gnoristini Gnoriste Meigen
Boletina Staeger
Coelosia Winnertz
Paratinia Mik
Deiedzickia Johannsen

Leiini Leia Meigen
Tetragoneura Winnertz
Ectrepesthoneura Ender-
lein

Docosia Winnertz
Anomalomyia Hutton

Cycloneurini Cycloneura Marshall
Procycloneura Edwards
Allactoneurini Allactoneura De Mei-
jere *
Mycetophilinae Exechini Allodia Winnertz

Brachypesa Winnertz
Exechia Winnertz
Mycetophilini Cordyla Meigen
Epicypta Winnertz
Mycetophila Meigen (in-
cluding Mycothera
Winnertz )
Opistholoba Mik
Phronia Winnertz
Sceptoma Winnertz

Manotinae Manota Williston *
Lygistorhininae Lygistorhina Skuse *
Family Sciaridae Sciara Meigen

Pseudosciara Schiner

Subfamily BoLIropHILINAE

The subfamily Bolitophilinae is represented by the two genera
Bolitophila (fig. 1) and Bolitophilella (fig. 2), which have been dis-
tinguished on the basis of the termination of Ry of Edwards. In
Bolitophila this vein ends in the costa, whereas in Bolitophilella it
ends in Ry... The value of this character is somewhat open to ques-
tion as far as its use to separate genera is concerned. From the thoracic
sclerites, only minor differences are apparent. The episternum of the
prothorax is more elongate in Bolitophila and the mesepimeron is
slightly broader. However, the structures are so similar that we would
not recognize the two genera as distinct.

Subfamily DiromylINAE

The subfamily Ditomyiinae is represented by Centrocnemis (fig. 5),
Symmerus (fig. 3), and Nervijuncta (fig. 4). On the basis of thoracic

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

2 BOLITOPAILELLA S.GENTROGNEMIS

3.SYMMERUS

6. DIADOCIDIA 9.CEROPLATUS
Fics. 1-9.—Genera of the family Mycetophilidae.

NO. 2 FUNGUS GNATS—SHAW AND SHAW 5

sclerites Symmerus and Nervijuncta seem closely related. Both
genera possess a dorsal projection from the katepisternum of the
mesothorax. In reality this structure represents a fusion of the pos-
terior portion of the anepisternite with the katepisternite. In Sym-
merus the mesepimeron is greatly reduced, with only the dorsal and
ventral portions remaining. In Nervijuncta this reduction has gone
even further, with only the dorsal portion remaining.

We have never seen a specimen of Calliceratomyia Lane, but
through the courtesy of John Lane we have a sketch of the thorax of
a specimen of this genus. Though lacking in certain details, the draw-
ing clearly indicates that Calliceratomyia, on the basis of thoracic
sclerites, is closely related to Nervijuncta and Symmerus.

We are uncertain whether the genus Centrocnemis is correctly
placed in the Ditomyiinae. Unfortunately, we were not able to obtain
specimens of Ditomyia for study. The only characteristic that Cen-
trocnemis, Symmerus, and Nervijuncta show in common is the well-
developed metapleura. Otherwise, from the standpoint of thoracic
sclerites there is not much similarity. Centrocnemis possesses a well-
developed and broad mesepimeron resembling that of Apemon. The
dorsal lobe of the katepisternum, as found in both Symmerus and
Nervijuncta, is lacking. From the standpoint of venation, Centro-
cnemus is more closely related to Mycetobia (Anisopodidae) than to
Symmerus or Nervijuncta. Until the opportunity to study Ditomyia
is presented we reserve judgment as to the ultimate systematic position
of Centrocnemis.

Subfamily DrapocmmINaE

The subfamily Diadocidinae is represented by the genus Diadocidia
(fig. 6). The position of Diadocidia is not entirely clear. From
the standpoint of venation, this genus must be considered as more
highly evolved than either the Bolitophilinae or the Ceroplatinae. The
radius is 2-branched and the stem of media is lacking. The affinities
of Diadocidia seem to be closer to the Bolitophilinae than to the Cero-
platinae. Both Bolitophila and Diadocidia exhibit a remnant of a
meron. In both genera the anepisternal cleft is not so pronounced as
it is in Palaeoplatyura. The structure of the prothorax is similar for
Bolitophila and Diadocidia.

Subfamily CrEROPLATINAE

The Ceroplatinae figured represent the seven genera Palaeoplatyura
(fig. 7), Proceroplatus (fig. 8), Ceroplatus (fig. 9), Platyura (fig.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

10), Apemon (fig. 11), Macrocera (fig. 12), and Fenderomyia (fig.
13). In addition we have examined a specimen determined by Ed-
wards as Platyroptilon miersi Westwood. This specimen lacked an-
tennae; the pleurotergites were not hairy as Tonnoir (1929) states
them to be in this genus; and there was no evidence of setae on the
anepisternite. There may therefore be some question as to the cor-
rect identity of the insect. In any case the specimen we examined
identified as Platyroptilon was very close to Ceroplatus on the basis of
thoracic sclerites.

If Apemon be excluded, it is fairly simple to consider the other
genera of the Ceroplatinae as having developed from a form similar
to Palaeoplatyura. All exhibit the reduction of the mesepimeron and
a dorsoventral compression of the thorax resulting in an apparent
shifting of the sclerites posteriorly to a more nearly horizontal posi-
tion. Such a shift becomes very prominent in certain genera of the
Mycetophilini.

Apemon, as indicated in an earlier paper (Shaw, 1948), is some-
what of an anomaly. On the basis of thoracic sclerites its affinities
seem closer to Centrocnemis, Symmerus, and Nervijuncta of the
Ditomyiinae. As in Centrocnemis, the mesepimeron is broad. The
cleft in the anepisternite is very distinct and indicates what has prob-
ably happened in both Symmerus and Nervijuncta where the katepi-
sternite seems to have a dorsal lobe extending to the wing basis. On
the basis of thoracic sclerites Apemon occupies an intermediate posi-
tion between the Ditomyiinae and Ceroplatinae; from the standpoint
of venation it seems to be intermediate between Palaeoplatyura and
Platyura. As in Palaeoplatyura the stem of media is distinct. How-
ever, as in Platyura the r-m cross vein is lost, apparently through the
fusion of a portion of the stem of media with Rs. Apemon also ex-
hibits the dorsoventral flattening of the thorax as also found in Pro-
ceroplatus and Platyura.

The systematic position of the Macrocerinae has been somewhat in
question. Some workers, including Lane and Coher, consider this sub-
family inseparable from the Ceroplatinae. We have two representa-
tives of the Macrocerinae, Macrocera (fig. 12) and Fenderomyia
(fig. 13). Macrocera on the basis of thoracic sclerites is intermediate
between Palaeoplatyura and Platyura. However, in Palaeoplatyura,
Ceroplatus, and Platyura there is a rather indistinct indication of a
meron in the mesothoracic leg. This is lacking in Macrocera, Fender-
omyia, and also in Proceroplatus. Macrocera, Proceroplatus, Platy-
ura, and Ceroplatus all exhibit one characteristic in common—the re-
duction of the lower portion of the epimeron of the mesothorax. This

NO. 2 FUNGUS GNATS—SHAW AND SHAW 7

15. PARATINIA
12.MACROCERA 18.NEURATELIA
Fics. 10-18.—Genera of the family Mycetophilidae.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

culminates in the condition shown in Fenderomyia where the pleuro-
tergite and the katepisternite touch each other, obliterating practically
all the lower half of the epimeron but the tip. Ceroplatus, Procero-
platus, Platyura, and Fenderomyia also possess another characteristic
in common—the mediotergite instead of being rounded, as in Palaeo-
platyura and Macrocera, becomes distinctly angulated. This might
result from a dorsoventral flattening of the thorax and is also indi-
cated in the position of the pleurotergies, which shift to a more hori-
zontal position. Our present opinion would be that the Macrocerinae
should be included in the Ceroplatinae.

Subfamily ScrioPHILINAE

The Sciophilinae are represented by six tribes: The Mycomyiini,
the Sciophilini, the Gnoristini, the Leiini, the Cycloneurini, and the
Allactoneurini.

Tribe MYCOMYIINI

The Mycomyiini are represented only by the genus Mycomyia
(fig. 14). Shaw (1948) indicated that Mycomyia might be an an-
nectant form between the Sciophilinae and the Mycetophilinae. Such
a belief is based on the structure of the pleura, and also, as pointed
out by Fisher, the male hypopygium indicates such a relationship.

Tribe SCIOPHILINI

Edwards (1925) states that the possession of macrotrichia on the
wing membrane is diagnostic of the tribe. The value of this character
for generic recognition may be open to question. He adds that where
the microtrichia have disappeared it may not always be easy to deter-
mine which set of hairs is present. Another characteristic of value
in delimiting this group is the possession of some hairs or bristles
on the postnotum (mediotergite). This characteristic is not common
to all genera.

The Sciophilini figured in this study include eight genera—Eudi-
crana (fig. 21), Neuratelia (fig. 18), Paratinia (fig. 15), Parvicellula
(fig. 16), Phthinia (fig. 19), Polylepta (fig. 17), Sciophila (fig. 22),
and Syntemna (fig. 20). Monoclona was examined but not figured.
Of the species studied, all but Paratinia, Syntemna, and Monoclona
have hairs or setae on the mediotergite. All but Paratinia have setae
on the pleurotergites. Concerning the latter genus, Edwards (1925)
states: “It does not seem to be very closely related to the other genera
of the Sciophilini but I include it here on account of macrotrichia on

NO. 2 FUNGUS GNATS—SHAW AND SHAW 9

2!1.EUDICRANA

24.BOLETINA 27.CYCLONEURA
Fics. 19-27,—Genera of the family Mycetophilidae.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

the apical half of the wing and the elongate abdomen with large
seventh segment. It may be related to Phthinia and in some respects
appears intermediate between that genus and Speolepta.”

On the basis of the thoracic sclerites we see no evidence to support
such a belief. The structure of epimeron, the indication of fusion
of the posterior part of the anepisternite with the epimeron, and the
absence of setae on both mediotergites and pleurotergites indicate
that the affinities of the genus are closer to those of certain of the
Gnoristini than to the Sciophilini. If the possession of macrotrichia
is considered as the important characteristic, then Paratinia must be
included with the Sciophilini. If so placed it would be closer to Par-
vicellula than to other members studied. Possibly too much value has
been placed on the presence or absence of macrotrichia on the wing
in the past. The genus Parvicellula, according to Tonnoir and Ed-
wards (1926), is peculiar to New Zealand. In venation it most closely
resembles Monoclona. On the basis of the shape of the anepisternite,
Monoclona resembles Mycomyia. In fact, Parvicellula, Monoclona,
and Mycomyia seem fairly closely related on the basis of thoracic
sclerites. Parvicellula is not too closely allied to other members of the
Sciophilini on the basis of thoracic sclerites. The anepisternite and
katepisternite are more nearly equal in size in Parvicellula, and in this
respect the genus is more closely related to Phthinia than to the other
genera of this tribe represented in this study. The shape of the mes-
epimeron differs in both genera. The course of the dorsal half of the
mesopleural suture in Parvicellula may indicate that the posterior
part of the anepisternite may be fused with the epimeron at times. If
the pleural suture were to extend in a straight line dorsally from the
juncture of the anepisternite and the katepisternite, then the epimeron
would have the structure as shown in Paratinia. The indication is
that the posterior portion of the anepisternite may be fused with
either the mesokatepisternite or the mesepimeron. The genus Poly-
lepta, as indicated by Shaw (1948), has thoracic sclerites resembling
those of Platyura except that the dorsoventral flattening, as indicated
in Platyura, is not so marked in Polylepta. The possession of setae on
the anepisternite, pleurotergite, and mediotergite is possibly evidence
that Parvicellula and Polylepta are related.

The structure of the mesepimeron indicates that Polylepta and
Tetragoneura (Leiini) were related. This may be only a superficial
resemblance. Tetragoneura does not possess setae on the anepisternite,
the pleurotergite, or the mediotergite. In the wing venation Polylepta
and Neuratelia are very similar except for loss of vein R, (Edwards)
in Neuratelia. From the standpoint of thoracic sclerites the arching of

NO. 2 FUNGUS GNATS—SHAW AND SHAW 1h

the posterior pronotum is common to both genera. In Neuratelia the
posterior portion of the anepisternite seems to be in the process of
fusing with the epimeron.

Phthinia resembles in some ways Polylepta. Like Polylepta it has
setae on the mediotergite and pleurotergites. The anepisternites in
both genera have fine hairs. Phthinia shows evidence of specializa-
tion in venation and also in the subequal anepisternite and kat-
episternite of the mesothorax. Edwards (1925) has indicated that
Phithinia may be related to Speolepta. However, the latter genus
lacks setae on the mediotergite and on the pleuroterga. Since we do
not have material to study the sclerites of Speolepta, we cannot state
whether these structures affirm Edwards’ beliefs.

The genus Syntemna seems to be somewhat closely related to Neu-
ratelia and Polylepta on the basis of thoracic sclerites. The epimeron
of Polylepta might result from the fusion of a part of the posterior
lobe of the anepisternite with the mesepimeron. The venation of
Syntemna does not appear especially close to that of either of these
genera. Edwards (1925) has indicated that Syntemna may be more
closely related to the Gnoristini, apparently considering that Dzied-
zickia and this genus were similar. The thoracic sclerites do not
indicate too close a relationship. The prothorax of Dziedzickia is
greatly modified. Apparently the posterior lobe of the anepisternite
has fused with the mesepimeron in Dzgiedzickia. This would indicate
that the latter genus was more specialized than Syntemna. This view
is not supported by venation. Edwards’ inclusion of certain species of
Syntemna in Dziedzickia on the absence of macrotrichia on the wing
might be questionable unless supported by other characteristics.

The genus Sciophila seems to be the most specialized of the genera
studied in this tribe. While it has maintained a relatively broad mes-
epimeron, the suture between the anepisternite and katepisternite of
the mesothorax is almost lost. This condition is found also in the
genus Anomalomyia (Leiini). Moreover, there is a reduction in the
size of the epimeron of the prothorax.

The genus Stenophragma, represented by Stenophragma nigricauda
Edwards, on the basis of thoracic structures seems very closely allied
to Sciophila.

On the basis of venation Monoclona is close to Sciophila except for
the unbranched cubitus and poor development of anal veins. Edwards
(1925) indicated that the macrotrichia of the wing are reflexed in
Monoclona and not decumbent as in Sciophila. The thoracic sclerites
do not indicate a close relationship of the two genera. The anepister-

12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

nite of Monoclona resembles that of Mycomyia more closely than it
does that of Sciophila.

The genus Eudicrana is closely related to Sciophila. .The suture
between the mesothoracic anepisternite and katepisternite is beginning
to disappear. The prothoracic epimeron shows a reduction in its width.
From the standpoint of venation, Eudicrana seems more primitive.
Only in the absence of the median ocellus is Eudicrana more spe-
cialized than Sciophila.

Tribe GNORISTINI

The tribe Gnoristini is represented by four genera—Boletina (fig.
24), Coelosia (fig. 26), Dziedzickia (fig. 25), and Gnoriste (fig. 23).
On the structure of the thoracic sclerites the group appears to be quite
homogeneous. All possess an epimeron that is broad dorsally and nar-
rowed ventrally. Gnoriste seems to be most primitive on the basis of
the structure of the epimeron. However, Guoriste shows evidence
of specialization in the possession of an elongate proboscis.

We believe that in all this group the posterior part of the anepi-
sternite of the mesothorax is in the process of being fused or has been
fused with the mesepimeron. This condition is most marked in
Deiedzickia.

Boletina and Deziedzickia differ from Gnoriste and Coelosia in the
possession of setae on pleurotergites. In regard to Coelosia, Edwards
(1925) removed Phthinia thoracia Winnertz and P. curta Johannsen
to this genus. Only future examination of specimens of these will
determine if the thoracic sclerites indicate the validity of this grouping.
Another feature shared in common by the four genera studied is the
position of a suture between the anepisternite and katepisternite of
the mesothorax. This suture does not extend in a horizontal line, as
in many genera, but dips posteriorly. Both Gnoriste and Boletina
show a reduction in the size of the propleura. They also exhibit a
tendency for the pronotum to be compressed and eventually to as-
sume an almost horizontal position, as shown in Dziedzickia.

The venation of the genera studied varies considerably. Thus
Dziedzickia is the only one of the genera studied that possesses the
upper branch of the radial sector (Ry of Edwards) ; in other respects
the venation resembles that of Syntemna of the Sciophilini. Bole-
tina and Gnoriste have a similar venation. Coelosia resembles the
genus Phthinia of the Sciophilini, differing primarily in the loss of Sco.

Johannsen (1911) has noted that certain species of Boletina,
Coelosia, and Gnoriste share the peculiarity of having one claw of
each foot of the male modified.

NO. 2 FUNGUS GNATS—SHAW AND SHAW 3

Tribe LEIINI

The tribe Leiini is represented by the genera Leia (fig. 33), Tetra-
goneura (fig. 30), Ectrepesthoneura (fig. 29), Docosia (fig. 31),
Anomalomyia (fig. 32). The genera Cycloneura and Procycloneura,
formerly placed in the Leiini, are considered to represent a distinct
tribe, the Cycloneurini.

All these genera lack setae on the mediotergite. Leia and Ano-
malomyia possess setae on the pleurotergites. Procycloneura and
Anomalomyia have lost the suture separating the mesothoracic an-
episternite from the katepisternite. The condition has been noted
already in Sciophila (Sciophilini). The significance of some of these
apparent relationships may be open to question.

Ectrepesthoneura and Tetragoneura are closely related. Some
slight differences in the shape of the pronotum, the mesothoracic
katepisterna, and epimera. These are probably of sufficient impor-
tance to justify the maintaining of the two genera as distinct. The
two genera differ in venation: Tetragoneura has Sc short and ending
free and Cu forking near the middle of the wing; Ectrepesthoneura
has Sc longer, ending in R, and Cu forking near the base of the wing.
Edwards (1925) calls attention to the forking of the cubitus near
the wing base, supporting the conclusion that this group of genera are
more or less related to the Sciarinae. On the basis of thoracic
sclerites, the similarity between Tetragoneura and Ectrepesthoneura
is not so evident as would be indicated from the venation. Docosia,
on the basis of thoracic sclerites, more closely resembles the Sciaridae.
However, the resemblance is more or less superficial. Sciara possesses
a midpleural pit, the division of the pronotum into an anterior and a
posterior division, and a precoxal bridge. All these characteristics
are lacking in Docosia. Docosia, on the basis of thoracic structures,
is not closely related to either Leia or Anomalomyia.

Edwards (1925) pointed out that Docosia in life much resembles
Sciara. He adds that Docosia also resembles Tetragoneura in appear-
ance and habits and concludes that the genus is not distantly related
to Tetragoneura. The shape of the epimeron, the pleurotergite, and
the course of its suture separating the mesothoracic anepisternite
from the katepisternite do not support the close relationship of Tetra-
goneura and Docosia. The absence of pleurotergal setae is common
to Docosia, Ectrepesthoneura, and Tetragoneura.

Anomalomyia, on the basis of venation, resembles rather closely
Rondaniella. It would not be too difficult to derive the shape of the
thoracic sclerites as found in Anomalomyia from those of Leia. If

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

28. PROCYCLONEURA

31.DOCOSIA

29, ECTREPESTHONEURA 32. ANOMALOMYIA

36. BRACHYPEZA

30. TETRAGONEURA
33.LEIA

Fics. 28-36.—Genera of the family Mycetophilidae.

NO. 2 FUNGUS GNATS—SHAW AND SHAW 15

the posterior lobe of the anepisternite were fused with the epimeron
as somewhat indicated in Leia, and if the suture between the anepi-
sternite and the katepisternite were completely lost, the two thoraces
would be quite similar. There are indications of loss of the suture
between the anepisternite and the katepisternite in Leia.

Cycloneura and Procycloneura might be placed in a tribe by them-
selves. Their inclusion in the Leiini seems questionable to us. In some
respects they seem closer to the Mycetophilinae. Both have Sc short
ending free. Procycloneura has the lateral ocelli touching the eye mar-
gin. They lack setae on the anepisternites, the pteropleurites, the pleu-
rotergites, and the mediotergites. However, there is some variation in
the distribution of setae in the representatives of the Mycetophilinae
studied. The coxae appear stouter and more compact as in Sceptonia
or Epicypta (Mycetophilinae). Until more material is made available
for study it may be well to maintain Cycloneura and Procycloneura
in the Sciophilinae. We do consider it valid to designate a new tribe—
the Cycloneurini—which we establish here for these two genera and
characterize as follows:

CYCLONEURINI, new tribe

Wing venation of the same general type as in Leia. Some species
of Cycloneura with a fusion of Cu, and the anal vein, resulting in the
formation of a closed cell. Thorax showing dorsoventral depression.
Coxae stout and compact. Metapleura reduced in size.

In those forms possessing a suture between the anepisternite and
the katepisternite, the thoracic sclerites are subequal in size. In this
respect, the two genera placed in the Cycloneurini differ from the
Leiini studied, which have the katepisterna larger than the anepis-
terna. We would expect Paracycloneura to fall in this tribe, but not
having seen specimens we cannot definitely place it here.

Of the two genera, Cycloneura, on the basis of thoracic sclerites,
appears the more primitive. The suture between the anepisternite
and the katepisternite of the mesothorax persists but is lost in Procy-
cloneura. Also the dorsoventral compression of the thorax is more
marked in the latter genus.

ALLACTONEURINI, new tribe

A specimen of Allactoneura argentosquamosa Enderlein, deter-
mined by Edwards, was made available for examination by Dr. Free-
man. Edwards (1925) erected a new subfamily, the Manotinae, to

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.) 287

include this genus and Manota. An examination of thoracic sclerites
indicates that Allactoneura is very similar to Procycloneura. The
prothoracic region is more modified, being more dorsally produced.
The pleurotergites are setose. The venation does not resemble either
of the genera included thus far in the Cycloneurini. Allactoneura in
our opinion is not closely allied to Manota but is more closely related
to the Cycloneurini. It does not seem to agree with the Cycloneurini
in venation or in chaetotaxy. Edwards (1925) called attention to the
shape of the head of this genus, reminding one of the Brachycera and
Cyclorrhapa, and also to the presence of scales on the thorax and
abdomen. We consider it best to erect the above-named new tribe,
the Allactoneurini, for this genus. It may be characterized as follows:
Wing venation—Sc long, ending in costa, Scz present. R 2-branched,
M 2-branched. Cu 2-branched with fork at base of wing. Prothorax
projecting dorsally into mesoscutum. Suture between anepisternite
and katepisternite of mesothorax lost. Legs robust. Thorax and
abdomen with scales. At present this tribe includes only one genus—
Allactoneura.

Subfamily LycistoRHININAE

This subfamily was proposed by Edwards for Lygistorhina Skuse,
including Probolaeus Williston and Palaeognoriste Meunier. Johann- |
sen (1911) included Probolaeus with the Mycetophilinae but stated,
“Tt is possible that this genus should be placed with the Sciarinae.”
Although lacking proper material for a detailed study of the thorax,
an examination of some slides indicates that the affinities of Lygisto-
rhina are with those of the Sciophilinae, possibly being closest to the
Gnoristini. However, the peculiar head structure, the elongate pro-
boscis wholly unlike that of other mycetophilids, and the venation
warrant the maintenance of a separate subfamily.

Subfamily MycreTropHILINAE

The subfamily Mycetophilinae has been divided into two tribes on
the following characteristics by Edwards (1925) :

Anepisternal bristles present, hind coxa usually lacks a basal seta. . Mycetophilini
Anepisternal and pteropleural bristles absent, hind ccxa usually with a
Strone Wasal wSebae-ius sitios Sisteeaeno crenartaiow Goh come cio ae Exechini

Tribe EXECHINI

The Exechini are represented by Allodia (fig. 35), Brachypeza
(fig. 36), and Exechia (fig. 34) in this study.

NO. 2 FUNGUS GNATS—SHAW AND SHAW M7,

Exechia and Allodia are closely allied. They are separated by the
position of the fork of Cu, which is beyond that of media in Exechia
and before the medial fork in Allodia. On the basis of thoracic
sclerites there are no significant differences between the two. This
substantiates the view of Edwards (1925).

Brachypeza, on the basis of thoracic sclerites, is not too closely
allied to either Allodia or Exechia. In fact, our material has a setose
anepisternite and on this basis would not be placed in the Exechini.
It possesses one strong and two weaker setae on the base of the hind
coxa. Thus it possesses characters of both the Exechini and the
Mycetophilini.

On the basis of venation Brachypeza appears closely related to
Rhymosia. The anepisternite in some specimens of Rhymosia we have
examined resembles more closely the anepisternite in Brachypeza than
it does that of Allodia. This sclerite in Brachypeza is roughly hex-
agonal in shape, as in some of the species of Rhymosia we have ex-
amined. Possibly this may be of value in distinguishing between
Allodia and Rhymosia. An investigation should be made of the two
genera Allodia and Rhymosia to determine whether both are valid
and also to reevaluate the systematic position of the species of the
two genera.

Tribe MYCETOPHILINI

The tribe Mycetophilini is represented by Cordyla (fig. 37), Ept-
cypta (fig. 42), Mycetophila (fig. 39), Mycothera (fig. 40), Opistho-
loba (fig. 41), Phronia (fig. 38), and Sceptoma (fig. 43). There
seems to be a logical division of these genera on the absence or pres-
ence of pteropleural (mesepimeral) setae. On this basis, Dynatosoma,
Cordyla, Trichonta, and Phronia form one group and the remain-
ing genera of the tribe Mycetophilini another. This division seems
to be supported by the structure of the thoracic sclerites. We have
not examined Dynatosoma as far as the thoracic sclerites are con-
cerned. From the standpoint of venation, this genus seems close
to Trichonta. In general appearance the genus resembles Mycetophila
but lacks mesepimeral bristles.

Edwards (1925, p. 587) has figured the thorax of Trichonta. While
somewhat lacking in detail, it indicates that Phronia and Trichonta
are closely allied. Both have the anepisternite more or less hexagonal
in shape; the structure of the epimera is very similar.

Cordyla seems to be the most specialized of this group. It shows
more marked dorsoventral compression of the thorax. Evidence of
specialization is also indicated by the enlarged second palpal segment

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

39.MYCETOPHILA 42.EPICYPTA 45. PSEUDOSGIARA

Fics. 37-45.—Genera of the families Mycetophilidae and Sciaridae.

NO. 2 FUNGUS GNATS—SHAW AND SHAW 19

and the reduction of number of segments of the antennae. There is
a difference in specialization between the males and females in regard
to this aspect, since in general the females have fewer segments.

The remaining Mycetophilini are all characterized by the possession
of mesepimeral setae. It would seem possible to subdivide further
the genera studied, Epicypta, Mycetophila, Mycothera, Opistholoba,
and Sceptonia. In all but Mycetophila and Mycothera the mesepi-
meron shows a tendency to occupy somewhat of a horizontal position.
This group of genera also shows a marked dorsoventral compression
of the thorax.

Mycetophila and Mycothera are very closely related. Edwards,
(1925) united Mycothera and Opistholoba with Mycetophila. In an
earlier paper, Shaw (1948) indicated his reasons for regarding
Opistholoba as a distinct genus. Though minor differences do exist in
the thoracic sclerites of Mycetophila and Mycothera, it would be dif-
ficult to define such differences in a way to justify the separation of
these two genera.

The genera Opistholoba, Epicypta, and Sceptoma form a closely
related group. Edwards (1925, p. 587) figured Delopsis, and this
genus would seem to culminate the tendencies shown in this series.
The modifications include—

1. A progressively increasing dorsoventral compression of the thorax.

2. The mesepimeron tends to become more nearly horizontal in position.

3. The prothorax pushes dorsally, thus forming a concave region in the
lateral margin of the mesoscutum.

These genera can be separated on the basis of characteristics in
the thoracic sclerites. Thus Epicypita has the mesepimeron widened
at the apex. In degree of dorsoventral compression, it is intermediate
between Opistholoba and Sceptonia. Both Sceptonia and Delopsis
have the anepisternite subrectangular in shape. Sceptonia lacks the
expanded apex of the mesepimeron; moreover the mediotergite is
guite pointed. The pronotum has pushed noticeably into the meso-
scutum. The genus Delopsis seems to be the most highly specialized
of the genera studied. The mesepimeron is greatly reduced ; the pro-
notum extends deeply into the margin of the mesoscutum.

Subfamily MANOTINAE

Edwards (1925) recognized a separate subfamily for Allactoneura
and Manota. As discussed elsewhere in this paper, Allactoneura ap-
pears to us much more closely related to the Cycloneurini of the
Sciophilinae. However, Allactoneura does not agree with our con-

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

cepts of the Cycloneurini, and therefore we have erected a new tribe
—Allactoneurini—for this genus.

A specimen of Manota defecta Williston was lent for examination.
Since the specimen could not be boiled, it was impossible to make a
detailed study. Sufficient details could be observed to substantiate the
belief that a separate subfamily should be recognized. The prothoracic
region is large, expanded, and somewhat shieldlike. No distinct
suture could be seen between the prothoracic pleura and tergum. The
katepisternum and epimeron of the mesothorax appears to be fused,
thus forming a large single sclerite. The head is unlike that of other
Mycetophilidae with which we are familiar. The maxillary palpi are
wholly different from those of any other genus of Mycetophilidae.

Family SCIARIDAE

Two genera of the related family Sciaridae are included. There
has been much discussion as to the systematic position of this group.
Edwards (1925) indicated their similarity in appearance to Tetra-
goneura and Docosia (Leiini). He held that the Leiini and Sciarinae
probably had common origin, but since some of the more primitive
genera of sciarids maintain macrotrichia, he did not consider that this
group evolved from Tetragoneura or its near relatives.

The similarities in venation between the two groups represent a
case of parallel development of a characteristic by two separate groups.
This phenomenon has been noted elsewhere by Crampton. It indicates
the need for use of many characteristics rather than one or two in
grouping genera.

Sciara (fig. 44), as indicated in an earlier paper (Shaw, 1948), is
sufficiently distinct from the true Mycetophilidae on the basis of
thoracic sclerites to warrant its being placed in a separate family.

Pseudosciara (fig. 45) was placed with the Mycetophilinae by
Johannsen (1909), who states: ‘“The form of the head and the course
of media remind one of Sciara, but the long coxae and position of Cu
show relationship of Mycetophilini.’ Edwards (1925) considered
this genus to be allied to the Leiini (Mycetophilinae). In 1932 he
placed the genus with the Sciarinae.

On the basis of thoracic sclerites it is evident that Pseudosciara is a
true sciarid and that any similarities of this genus with the Mycet-
ophilinae are more apparent than real. Both Sciara and Pseudosciara
possess a distinct precoxal bridge, which is lacking in all the genera
of the Mycetophilidae we have studied. The shape of the other

NO. 2 FUNGUS GNATS—SHAW AND SHAW 21

sclerites is similar in both genera, with Pseudosciara appearing more
specialized. Both possess a midpleural pit.

SUMMARY

The pleural sclerites of 45 genera of fungus gnats are herein illus-
trated. A discussion of the relationships of these and certain other
genera is presented.

On the basis of the structure of the thoracic pleura, the genus
Bolitophilella is not considered as distinct from Bolitophila. The
genus Calliceratomyia, placed in the Ceroplatinae by Lane, is con-
sidered to be more closely related to Nervijuncta and Symmerus
(Ditomyiinae) than to the genera of the Ceroplatinae we have studied.
The systematic position of Centrocnemis is somewhat uncertain at
present. The genus Apemon appears to be intermediate between the
Ditomyiinae and the Ceroplatinae. It is logical to include Macrocera
and Fenderomyia in the Ceroplatinae. The genus Paratinia seems to
be more closely allied to the Gnoristini than to the Sciophilini, where
it had been placed by Edwards (1925). The genera Parvicellula,
Monoclona, and Mycomyia are fairly closely related and are apparently
intermediate between the Sciophilinae and the Mycetophilinae. Since
Cycloneura and Procycloneura do not appear closely related to the
other genera of the Leiini in which they were placed on the basis of
wing venation, a new tribe—the Cycloneurini—is proposed for these
two genera and probably for Paracycloneura. The genus Allactoneura
is removed from the Manotinae and placed in a new tribe—the Allac-
toneurini. The affinities of Lygistorhina seem to be closest to certain
of the Gnoristini, but we consider that a separate subfamily should be
recognized for Lygistorhina. The genus Mycothera does not appear
to be distinct from Mycetophila. However, Opistholoba seems to be
sufficiently distinct to warrant its maintenance as a separate genus.
The genus Manota is so different from the other genera that a sepa-
rate subfamily should be recognized for it.

Two genera (Sciara and Pseudosciara) of the related family
Sciaridae are figured. Pseudosciara has been variously grouped by
earlier investigators, some of whom have placed the genus in the
Mycetophilinae. The thoracic sclerites indicate that it should be
included with the other Sciaridae.

CONCLUSIONS

As suggested in an earlier paper (Shaw, 1948), the shape of the
pleural sclerites is of value as a means to indicate phylogenetic rela-

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Ti7

tionships of the Mycetophilidae. Future investigators would do well
to include these structures in describing new genera and species.

ABBREVIATIONS USED ON FIGURES

AES, Anepisternum of mesothorax.
APN, Anterior pronotum.

EM,, Prothoracic epimeron.

EM,, Mesothoracic epimeron.

EM,, Metathoracic epimeron.

ES,, Prothoracic episternum.

ES,, Metathoracic episternum.
KES, Katepisternum of mesothorax.
MP, Midpleural pit.

MT, Mediotergite.

PLT, Pleurotergite.

PPN, Posterior pronotum—pronotal scutellum.
PSc, Prescutum of mesonotum.

Sc, Scutum of mesonotum.

ScT, Scutellum of mesonotum.

REFERENCES
CRAMPTON, G. C.
1925. A phylogenetic study of the thoracic sclerites of the non-tipuloid
nematocerous Diptera. Ann. Ent. Soc. Amer., vol. 18, pp. 49-74,
pls. 3-7.
1942. The external morphology of the Diptera. Diptera of Connecticut,
fasc. I, pp. 10-165, figs. I-14.

Epwarps, F. W.
1925. British fungus gnats. Trans. Ent. Soc. London, 1924, pp. 505-670,
pls. 49-61.
1932. New Brazilian Mycetophilidae. Rev. Ent., vol. 2, pp. 138-149, figs.
1-6.

JOHANNSEN, O. A.
1909. Mycetophilidae. Genera insectorum, fasc. 93, pp. 1-140, pls. 1-7.

1909. The fungus gnats of North America: Pt. I. Maine Agr. Exp. Stat.

Bull. 172, pp. 209-276, 3 pls.
1910. The fungus gnats of North America: Pt. I]. Maine Agr. Exp. Stat.
Bull. 180, pp. 125-192, 4 pls.
19o1t. The fungus gnats of North America: Pt. III]. Maine Agr. Exp.
Stat. Bull. 196, pp. 249-328, 5 pls.
1912. The fungus gnats of North America: Pt. IV. Maine Agr. Exp.
Stat. Bull. 200, pp. 57-146, 6 pls.
LAnprock, K.
1940. Pilzmucken oder Fungworidae. Tierwelt Deutschlands, vol. 38, pp.
1-166, figs. 1-338.
SHaw, F. R.
1935. Notes on the Mycetophilidae with descriptions of new species. Psyche,
vol. 42, pp. 84-91, figs. 1-5.

NO. 2 FUNGUS GNATS—SHAW AND SHAW 23

1948. A contribution to the phylogeny of the Mycetophilidae. Ann. Ent.
Soc. Amer., vol. 41, pp. 189-199, pls. 1-3.
Tonnorr, A. L.
1929. Australian Mycetophilidae. Proc. Linn. Soc. New South Wales,
vol. 54, pp. 584-614, figs. 1-7, pls. 22, 23.
Tonnorr, A. L., and Epwarps, F. W.
1926. New Zealand fungus gnats. Trans. New Zealand Inst., vol. 57, pp.
747-878, pls. 58-80.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 4

A REVISED CLASSIFICATION FOR
ins BIRDS OF THE WORLD

BY
ALEXANDER WETMORE

Secretary, Smithsonian Institution

S200 00ece?

(Pustication 4057)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 1], 1951

The Lord Baltimore Press

BALTIMORE, MD., U. 8 A.

Peake ViIskD CLASSIFICATION, FOR (iiE BIRDS
OF ThE WORLD

By ALEXANDER WETMORE

Secretary, Smithsonian Institution

Since the revision of this classification published in 1940? detailed
studies by the increasing numbers of competent investigators in avian
anatomy have added greatly to our knowledge of a number of groups
of birds. These additional data have brought important changes in
our understanding that in a number of instances require alteration in
time-honored arrangements in classification, as well as the inclusion
of some additional families. A few of these were covered in an edition
issued in mimeographed form on November 20, 1948. The present
revision includes this material and much in addition, based on the au-
thor’s review of the work of others and on his own continuing studies
in this field. His consideration necessarily has included fossil as well
as living birds, since only through an understanding of what is known
of extinct forms can we arrive at a logical grouping of the species
that naturalists have seen in the living state. The changes from the
author’s earlier arrangement are discussed in the following paragraphs.

Addition of a separate family, Archaeornithidae, for the fossil
Archaeornis siemensi, reflects the evident fact that our two most
ancient fossil birds, Archaeopteryx and Archaeornis, are not so closely
related as their earlier union in one family proposed. The characters
marking the two have been under dispute in literature, mainly be-
tween Petronievics and von Nopsca. Lambrecht? has analyzed the
data, finding 10 points of difference that hold. Though some of these
appear trivial, enough are of sufficient weight from a taxonomic stand-
point to warrant family status. Neither the suggestion that what we
now recognize as Archaeornis is merely a young individual of Archae-
opteryx lithographica, so that the differences seen are those of im-
maturity, nor the more extreme view that Archaeopteryx and Archae-

1 Wetmore, A., A systematic classification for the birds of the world. Smith-

sonian Misc. Coll., vol. 99, No. 7, Oct. 10, 1940, pp. I-II.
2 Handbuch der Palaeornithologie, 1933, p. 86.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 4

Pa SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

ornis have given rise respectively to our flightless and our flying birds
in separate lines of descent is supported by the facts.

In the Neornithes one important result has been the arrival at a more
even continuity of characters that has led to the reduction of super-
orders recognized in the subclass from four to three, through the com-
bination under Neognathae of the orders formerly separated as the
Palaeognathae. For years I have felt that recognition of the Palae-
ognathae, as a separate group apart from other birds, on the basis
of a supposed peculiarity in the palate, stood on very flimsy ground.
The recent studies of McDowell * demonstrate that the structure of
the palaeognathous palate, in which the palatine and pterygoid bones
are articulated by a squamous suture, is variable from order to order,
and that in fact the details of this union vary considerably in the dif-
ferent groups. For example, McDowell points out that in Dromiceius
the palatine and pterygoid are not in contact, while in a number of
families placed in the Neognathae, as in the Anatidae, to name only
one, the two bones are in articulation. As there is no clear-cut sepa-
ration, the former Palaeognathae must be combined with the Neog-
nathae.

The question of the weight to be given the peculiarities of uniform
pterylosis, extreme specialization of the wing as a flipper for subma-
rine progression, and incomplete fusion in the metatarsal elements, as
well as such other details as erect posture in standing and walking and
the anatomical adjustments involved, found in the penguins, is one
that has merited careful review. It seems reasonable after this ex-
amination to retain the Impennes as a superorder, at least until we
have further evidence through fossils as to their line of evolution.
It is necessary, however, to remove the fossil family Cladornithidae,
since Simpson* has found that the two genera Cruschedula and
Cladornis placed in this family have no apparent relationship to the
Sphenisciformes. These two, described by Ameghino from the De-
seado formation of Patagonia, now placed in the Oligocene, are based
on fragmentary, considerably flattened metatarsi. The descriptions
and figures that have appeared thus far are not sufficiently definite to
demonstrate clearly characters of importance in classification. How-
ever, from what we now know these ancient birds cannot be considered
as ancestral penguins of terrestrial habit, as has been supposed. The
only suggestion that has come to me is that possibly they may belong

3’ The bony palate of birds. Part 1, the Palaeognathae. Auk, vol. 65, Oct.
1948, pp. 520-549, 6 figs.

* Simpson, George Gaylord, Fossil penguins. Bull. Amer. Mus. Nat. Hist.,
vol. 87, art. 1, 1046, pp. I-99, 33 figs.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE 3

in the order Pelecaniformes, in which I have placed the family tenta-
tively in the suborder Odontopteryges, where it is located with two
others of almost equally uncertain status. This allocation is wholly
tentative and is no indication of belief in close relationship in the
three diverse groups there assembled.

The family Eleutherornithidae is introduced for the fossil Eleu-
therornis helveticus Schaub, from the Eocene of Switzerland, de-
scribed from a fairly well preserved pelvis. Apparently this is repre-
sentative of an ancestral group from which the living ostriches have
come. Its greatest importance is found in its indication of relationship
with carinate groups though of unquestioned ratite stock. It is thus
important as definite indication that the struthious birds have come
from flying ancestors, and not from some distinct cursorial line that
always has been flightless, as some have contended.

The family segregation in the order Procellariiformes has been
oversimplified in some recent considerations, probably through mis-
understanding of the group characters produced by anatomical studies,
possibly also through somewhat confusing names that have been ap-
plied to familial and generic categories. The Diomedeidae and Pele-
canoididae have been accepted without apparent question, but the
remaining species have been combined by some under a single family
name. Lowe,® however, has shown that the genera included in the
Hydrobatidae have a simplified condition in the quadrato-tympanic
region of the skull in which the opening of the upper tympanic recess
is small and is so located that it separates the squamosal and opisthotic
facets. In addition, the posterior border of the sternum is truncated
and entire, and basipterygoids are absent or are represented only by
small spines. In the Procellariidae, on the other hand, the foramen
of the upper tympanic recess is greatly enlarged and lies anterior to
the two facets for the quadrate, which are joined by a bridge of bone ;
the posterior border of the sternum is notched; and basipterygoid
processes are present. These constitute distinctive characters at the
family level.

In the arrangement of suborders in the order Pelecaniformes we en-
counter in marked degree the standard difficulty of logical placement
in linear alignment of groups that really stand in three-dimensional
relationship. Lanham ° has made a summary of the major anatomical
characters of the group in which he points out the differences that
set off the Phaéthontes and the Fregatae from the Pelecani. There

5 Proc. Zool. Soc. London, 1925 (Jan. 14, 1926), pp. 1436-1443.
6 Auk, vol. 64, 1947, pp. 65-70.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

is no question that the first two carry primitive characters, which may
be presumed to be similar to those found in an ancient ancestral stock,
since in these resemblances they are more like other types of birds,
notably the Procellariiformes. From this style the other families of
the Pelecani have become widely divergent. Although the tropic-birds
and the frigate-birds both have retained a part of what may be re-
garded as a basic pattern, they are so widely divergent in other re-
spects that it seems more reasonable to relate them individually as
branches from the common stock rather than to combine the two on
one line, separate from the Pelecani. The Phaethontes possibly may
have separated earlier than the Frigatae. Among interesting differ-
ences other than those of internal anatomy, it may be noted that the
tropic-birds have the young covered with down at birth and that the
adults possess series of air cells under the skin on the forepart of the
body like those found in pelicans and boobies. The frigate-birds have
the young naked at hatching, and the emphysematous condition is
mainly lacking. In view of this discussion I prefer to continue to
align these groups on either side of the Pelecani.

Though there is no question that the cormorants and snake-birds
are closely allied, they differ in such degree that they should be re-
tained in separate family status. The snake-birds are marked by a pe-
culiar conformation of the cervical vertebrae through which the beak
becomes a triggered spear in feeding. The bridge of Donitz on the
ninth vertebra is an important part of this arrangement. The stomach
also is unusual in possessing a curious pyloric lobe, lined with a mat
of hairlike processes. And there is only one carotid artery.

The Odontopteryges, as has been noted above, are of highly doubtful
status and require further study.

The family Cochleariidae, which some wish to unite with the Ardei-
dae, is marked externally by the strangely expanded bill, which is not
only broad, but has the gonys remarkably shortened. Internally the
pattern set by the broadened premaxillae is reflected in correspond-
ingly widened palatines, which in addition have a curious flaring ex-
pansion of the lateral margin. The lachrymals are greatly reduced in
size, and there are other minor peculiarities. Ridgway” lists four
powder-down tracts for Cochlearius, a larger number than the two
or three pairs that he found in the Ardeidae. This, however, needs
further checking, since there is some uncertainty as to the correctness
of these figures. There is no question that the boat-bills are closely
related to the herons, but the greatly modified forepart of the skull

7 Studies of the American Herodiones. Bull. U. S. Geol. Geogr. Surv. Terr.,
vol. 4, No. 1, 1878, p. 220.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—WETMORE 5

seems sufficient to warrant separation in a distinct family. Herons,
with riflelike precision, habitually spear or seize their prey, while the
boat-bill has the advantage of a broadened scoop. Possibly this is more
efficient in shallow waters, particularly in nocturnal feeding.

In view of the fact that the structural characters of the Balaenicipit-
idae have been summarized clearly by Stresemann,® it seems strange
that the status of this family has been a matter of question. The
single species shows affinity both with storks and with herons, in ad-
dition to outstanding peculiarities of its own.

The flamingos, which show affinity with both Ciconiiformes and An-
seriformes, have been placed by Stresemann and others in a sepa-
rate order, but they seem best allocated as a suborder of the first-
named group.

The superfamily Neocathartoidea, and family Neocathartidae, for
the curious vulture Neocathartes grallator (Wetmore),® discovered
recently in the Upper Eocene fossil beds of Wyoming, introduce a
new element in our known avifauna in the form of a small-winged,
strong-legged vulture that evidently was terrestrial with limited
powers of flight. It had about the same relation to the other Ameri-
can vultures that the secretarybird has to the hawks and falcons.
Its inclusion also requires a separate superfamily, the Cathartoidea,
for the previously known cathartine families.

_ The Numididae, which have been placed by some as a subfamily of

the Phasianidae, differ in completely lacking the tuberosity or plate
on the inner side of the second metacarpal that is so prominent in
pheasants and grouse. The Tetraonidae, in contrast with the Pha-
sianidae, have the pelvis relatively much broader and different in
proportion, and the tarsus relatively shorter in relation to the length
of the tibiotarsus. With these differences in mind it seems reasonable
to retain the three groups in family status, at least until more detailed
knowledge of their anatomy as a whole warrants change.

In the Turnices the two genera of bustard-quails, Turnix and
Ortyxelus, have no hind toe, the wing is eutaxic, only the left carotid
is present, and the eggs are rounded oval. The plain-wanderer of
Australia, Pedionomus, has a small hind toe, the wing is diastataxic,
right and left carotids are found, and the large eggs are pyriform.
It seems desirable to continue these as separate families, rather than

8 Aves, in Kiikenthal and Krumbach, Handbuch der Zoologie, vol. 7, pt. 2, sect.
8, 1934, p. 809.

9 For the substitution of Neocathartes for Eocathartes Wetmore, 1944, pre-
occupied by Eocathartes Lambrecht, 1935, see Auk, vol. 67, 1950, p. 235.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

as subfamilies of one group, an arrangement that Stresemann 7° has
accepted.

In the suborder Phororhaci the family Psilopteridae is to be added
for the South American fossils Psilopterus and Smiliornis, from the
studies of Patterson.*t Another group of doubtful position that may
be placed tentatively in this suborder, at least until it is better known,
is the family Cunampaiidae, for the fossil Cunampaia simplex, named
by Rusconi 7? from the Oligocene of western Argentina.

In the Charadriiformes the main disagreements of the present day
are found in the superfamily Charadrioidea and the suborder Lari,
in which the groups have been regarded by some as of family value
and by others have been allocated the rank of subfamilies. The var-
ious structural studies that have been made have not been complete
from a taxonomic point of view except for part of the species, and
the conclusions derived from the data available have been in the main
more philosophical than concrete. The picture therefore still remains
confused.

In view of the diverse specializations that are apparent, and the
obvious long evolutionary history, it appears better to me to continue
to acknowledge the main segregations as families, at least until the
subjects involved have been more thoroughly investigated. A family,
Rhegminornithidae, is added for the fossil Rhegminornis calobates
Wetmore, described from the Lower Miocene of Florida. This was
as large as a medium-sized curlew, of peculiar form as regards the
foot, the only part of the skeleton known, which shows certain char-
acters that seems to point toward the jacanas, though the bird is to
be placed in the Charadrioidea.

In the Lari the terns and the gulls are regarded as one family,
though there are some reasons that make further examination of this
treatment desirable. The Rynchopidae differ decidedly in absence of
the ambiens and the biceps slip, and in the completely different form of
the bill. The iris opens and closes vertically in slitlike form, a condi-
tion that I have not seen in any other bird.?°

The Stercorariidae possess a 2-notched sternum, large caeca, a cere,
and a complex rhamphotheca. In the Laridae ambiens and biceps slip
are present, the sternum is 4-notched, there is no cere, and the rham-
photheca is simple in form.

10 Loc. cit., p. 760.

11 Field Mus. Nat. Hist., geol. ser., vol. 8, No. 8, Oct. 31, 1941, pp. 52-53.

12 Bol. Pal. Buenos Aires, No. 21, May 2, 1946 (p. 1).

13 Wetmore, A., A note on the eye of the black skimmer (Rynchops nigra).
Proc. Biol. Soc. Washington, vol. 32, Dec. 31, 1919, p. 105.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—WETMORE ai

Old World ornithologists in the main have regarded the owls as
belonging to a single family, but while all are deceivingly similar in
general aspect, the decided differences found in the barn-owl group
merit family recognition. Ridgway ** years ago summarized the con-
siderable structural characters separating the Tytonidae and the Strig-
idae. It is necessary here only to point out the more outstanding
structural differences of the Tytonidae in the lack of the manubrium,
and the different form of the posterior margin of the sternum, which
is entire or has two shallow notches, the union of the furculum with
the carina sterni, the straight outline of the palatines, and the ventral
pteryla where the outer branch joins posteriorly to the main tract.
The Strigidae possess a manubrium, the sternum is 4-notched, the
furculum is separate, the palatines are greatly expanded posteriorly,
and the posterior end of the ventral pteryla does not join the main
tract at the posterior end.

Lucas *° long ago demonstrated the differences between the true
swifts and the crested swifts, though his work seems latterly to have
been largely overlooked in view of the recent inclusion of the two in
one group, as by Stresemann and by Mayr and Amadon. The skull
in the Hemiprocnidae is quite distinct in the general form of the
cranium and in the development of the nasals, vomer, and palatines,
while the hypotarsus has a tendinal foramen (like that found in hum-
mingbirds), and the plantar tendons have the flexor longus hallucis
connected with the branch of the flexor perforans digitorum, which
extends to the fourth digit. Coupled with this there may be noted the
curious nest, which, fastened to the side of a branch, is barely large
enough to contain one egg, and the further fact that these birds perch
regularly on branches and twigs in trees.

As Apus Scopoli, published in 1777, is recognized now in place of
Micropus Meyer and Wolf, 1810, for the type genus of the swifts,
the terms in the classification change to order Apodiformes, suborder
Apodi, and family Apodidae, which replace the former terms Mi-
cropodiformes, Micropodi, and Micropodidae, respectively.

The proposal of Mayr and Amadon ** to include the rollers in one
family, the Coraciidae, with three subfamilies, goes back to the ar-
rangement of Dresser in his monograph of the group.’? Sclater,1®

14. S. Nat. Mus. Bull. 50, pt. 6, 1914, p. 508.

15 Auk, vol. 6, 1889, pp. 8-13; vol. 12, 1895, pp. 155-157.

16 A classification of recent birds. Amer. Mus. Nov., No. 1496, Apr. 2, 1951,
PP. 9, 35.

17 A monograph of the Coraciidae, or family of the rollers. London, 1893,
Pp. i-xx, I-III, 27 plates.

18 Proc. Zool. Soc. London, 1865, pp. 682-688, 8 figs.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

however, many years ago, pointed out the pelvic powder-down tracts,
the small manubrium and other peculiarities of Leptosoma, and set
it apart in a distinct family. The anatomy of the syrinx and feet was
further elaborated by Forbes.1® The family Leptosomatidae there-
fore should be recognized.

The ground rollers, Brachypteracias, Atelornis, and Uratelornis,
usually have been included as a subfamily of the Leptosomatidae, but
Stresemann ?° places them in a separate family, the Brachypteraciidae.
There seems to be reason for this in their general appearance, though
their anatomy is not well known. Brachypteracias, in its skeleton,
differs from Coracias and Eurystomus in the much greater depth
of the outer notch on the posterior border of the sternum, in the much
broader and stronger pelvis, the heavier femur, and the greater cur-
vature of the shaft and reduction of the crista superior of the humerus.
I have not seen the skull. The habit of life is markedly different. AI-
though anatomical material of the other genera is not presently avail-
able, it seems reasonable to accept Stresemann’s proposal. These
peculiar birds certainly are not closely allied to Leptosoma.

Lack of information on the anatomy of the wood-hoopoes must be
the reason for the recent nonrecognition of the Phoeniculidae as a
family separate from the Upupidae, since the two are quite distinct and
have been so recognized for many years. The external differences
are readily apparent. In the skeleton in Phoeniculus (of which I have
seen several examples) the posterior part of the nasal area is ossified,
there being only a small, narrow, elongated nasal opening ; the ecteth-
moid is much reduced; the anterior end of the pterygoids is broadly
expanded ; the sphenoidal rostrum is swollen at the anterior end, where
the expanded ends of the pterygoids join it; the quadrates are de-
cidedly larger ; the keel of the sternum is greatly reduced, being only
half as high as in Upupa; the furculum is broader ; the pelvis is nar-
rowed, and considerably elongated posterior to the acetabulum, with
the ischio-pubic fenestra greatly enlarged; and the tarsus is heavier
and broader, with two definite fenestra below the head. There are
other minor details. In all of the above the characters of Upupa are
directly opposite. The two groups appear to me to be sharply set off
as distinct families.

The Passeriformes, with more living species than all the other

19 Proc. Zool. Soc. London, 1880, pp. 465-475, 5 figs.
20 Loc, cit., p. 820.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE 9

orders combined, and far fewer fossil forms discovered to date, pre-
sent many difficult problems in their logical arrangement. The major
groups are clear, whether we rank them as suborders or superfamilies
being a matter of opinion. But the limits of numerous families con-
tained in these larger categories are uncertain. The internal anatomy
is known fully for so few kinds that details of difference are poorly
understood. The superficial resemblances, on the other hand, are so
obvious in many cases that there is much confusion. Under the cir-
cumstances it continues to seem appropriate to me to accept the family
grouping that has been current for many years, except in those cases
where detailed studies clearly indicate change. There is much sup-
position in these matters, that has led to various proposals for combina-
tion, some part of which undoubtedly will prove correct. It is equally
probable that a part, possibly a considerably larger part, may prove
to be unfounded when details are more clearly known. If change is
accepted under these circumstances it may prove unwarranted, neces-
sitating further change, perhaps a return to the original supposition.
Since this can only prove confusing I prefer the conservative course.
In the remarks that follow I will discuss only a few matters on which
I have more or less concrete ideas.

In the superfamily Furnarioidea, Von Ihering *! unites the Fur-
nariidae and the Dendrocolaptidae, since he is unable to separate two
groups on the basis of the form of the posterior border of the nasal
opening. The variation that he shows seems quite true, but there are
numbers of other points of supposed difference concerned in the os-
teology and other structural details, and therefore his suggestion is
far from established. Pycraft,?? though seemingly uncertain in the
beginning, finally retained the two families. It may prove that some
genera are wrongfully allocated at present between the two groups,
and that their shifting, when we have sufficient information, will clear
our understanding.

In the Tyrannoidea, the family Oxyruncidae is known through ex-
ternal characters that seem to warrant separation. If the sharpbills
have other affinities it is doubtful that these are within the Tyrannidae,
where some have placed them.

In the family Cracticidae, recognized by Australian ornithologists,
the skull according to Pycraft ** (mainly from examination of Gym-
norhina) has the zygomatic process of the squamosal bifurcate, the
postorbital process large, the orbitosphenoid ossified, the interorbital

21 Auk, vol. 32, 1915, pp. 145-153, pls. 11-12.
22 Proc. Zool. Soc. London, 1906, pp. 133-159, figs. 49-52.
28 Proc. Zool. Soc. London, 1907, pp. 355-365.

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL ay

septum with a single opening, the prefrontals unusually large, and the
form of the palate peculiar. In his phylogenetic tree Pycraft places
the group on a common stem with the Artamidae, and not far from
the Paradiseidae. His account is mainly descriptive and difficult to
summarize in concrete form.

The family Grallinidae is likewise recognized officially by Austra-
lian ornithologists for Grallina cyanoleuca, the magpie-lark. The
principal study of the osteology is that of Shufeldt,?* but the account
is mainly descriptive and without definite conclusion. Amadon ** re-
cently has placed Corcorax and Struthidea here tentatively, though
this seems subject to further proof.

Stonor 7° has outlined excellent reasons for recognition of the
Ptilonorhynchidae, finding that they differ from Paradiseidae, with
which they have been united, in having an apterium in the center of
the dorsal feather tract, the tip of the vomer convex, larger, more de-
veloped maxillo-palatines, the margin of the palatines angular, smaller
ectethmoid, much larger lachrymal, and slender, greatly elongated
orbital ramus of the quadrate. The genera Loria and Loboparadisea,
usually included here, he transfers to the Paradiseidae. His conclu-
sion is that ‘‘the Ptilonorhynchidae constitute a singularly complete
and isolated family of the acromyodian passerine birds and show no
special relationship to any other, being sharply marked off by the
structure of the skull, the colour-pattern, and the bower-building
habit.”

Oberholser 77 has set up a distinct family Irenidae for the fairy
bluebirds (Jrena), and Delacour ** a family Aegithinidae for the
leafbirds, which would cover Irena, Aegithina, and Chloropsis. Inas-
much as the internal anatomy of these seems as yet unknown, I have
not included such a family, pending further information.

The proper allocation of the genus Chamaea for the wren-tits, at
present accepted by the A. O. U. Committee on Classification and No-
menclature as a separate family, the Chamaeidae, is one of consider-
able uncertainty. Delacour ?° has suggested that they be located in the

24 Emu, vol. 23, July 1923, pp. 16-10, pl. 6.

25 Emu, vol. 50, Oct. 1950, pp. 123-127.

26 Proc. Zool. Soc. London, vol. 107, ser. B, pt. 3, Sept. 1037, pp. 475-490,
figs. 1-9. It should be noted that the names on figures 6 and 8 have been trans-
posed, figure 6 being Semioptera wallacei, and figure 8 Amblyornis subalaris,
and not the reverse as printed on pp. 481 and 483.

27 Journ. Washington Acad. Sci., vol. 7, Oct. 19, 1917, pp. 537-541.

28 Zoologica, vol. 31, 1946, p. 3.

29 L’Oiseaux, vol. 16, 1946, pp. 18, 25, 35.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE II

family Timaliidae in a special subfamily in which he includes also
such diverse genera as Chrysomma (Moupinia), Panurus, Cono-
stoma, and Paradoxornis (combining under this name Suthora, Psit-
tiparus, Neosuthora, and Cholornis). This is an obviously hetero-
geneous assemblance, in which Chamaea has slight resemblances to the
first only. From Moupinia poecilotis (placed in Chrysomma by Dela-
cour) the wren-tit differs definitely in weaker, less arched bill and in
differently proportioned feet. It has no close resemblance to any of
the others that are mentioned. Although the relationships of Chamaea
are obviously uncertain, it is retained as a family pending other
information.

The recent suggestions for the union of the Bombycillidae, Ptil-
ogonatidae, and the Dulidae in one family are not substantiated by
examination of the skeleton. Dulus, the palm-chat, is widely differ-
ent from the other two, a structural distinction that is further em-
phasized by its curious communal nesting habits. The first two seem
more closely related but are separated clearly by characters found
in the ectethmoid region of the skull, and in the manubrium, to men-
tion only two points that are easily apparent. Delacour and Amadon °°
consider Hypocolius closely allied to Ptilogonys.

While Zimmer ** believes that the family Vireolaniidae should be
included in the Vireonidae, separate family rank in my opinion is
definitely justified. In addition to characters assigned by Pycraft*?
for the shrike-vireos I have found recently that in the pterylosis the
dorsal tract on the lower back is forked, the arms being broad at the
ends, and separated from the narrowed line that continues onto the
caudal area. This is completely different from the usual rhomboid
found in the vireos, and may indicate that the family eventually
should be removed from the vicinity of the Vireonidae.

The family characters of the pepper-shrikes, likewise outlined by
Pycraft in the reference given above, are easily apparent on examina-
tion of the skeleton.

The family Callaeidae has been separated by Stonor ** on the
weakened keel of the sternum, the great development of the lower
limb, coupled with reduced powers of flight, and the presence of a
mouth wattle, for three peculiar genera, Callaeus, Heterolocha, and
Philesturnus of New Zealand.

3° This, 1949, pp. 427-426, pl. 19.

31 Amer. Mus. Nov., No. 1160, Jan. 30, 1942, p. I0.
32 Proc. Zool. Soc. London, 1907, pp. 352-379.

33 This, 1942, pp. 1-18, figs. 1-10, tables 1-3.

{2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOUS 7,

The former family Melithreptidae becomes the family Melipha-
gidae, since the name of the type genus is now accepted as Meliphaga
Lewin, 1808.

In a similar way the family Compsothlypidae for the wood warblers
becomes the family Parulidae, since the former Compsothlypis
Cabanis, 1851, is replaced by the older Parula Bonaparte, described
in 1838.

The order of arrangement in the Passeriformes is in part neces-
sarily arbitrary, through the easily perceptible and often remarked
fact that we are under necessity of listing groups in linear order in a
two-dimensional alignment when actually they stand in three-dimen-
sional relationship to one another. (There is, further, another element
that may be regarded almost as a fourth dimension, in some of the
extinct groups known only as fossils that have no close relatives alive
today.) The sequence that I have adopted is the one that best repre-
sents my present understanding, based on personal studies that now
have extended over a period of nearly 50 years. I will repeat what I
have said elsewhere, that I have placed the Fringillidae at the end of
the list, because of my feeling that this group is the modern expres-
sion of a main core or stem that through the earlier Tertiary periods
has given rise to more specialized assemblages that we now recognize
as distinct families. Further specialization is apparent in some parts
of the existing fringilline assemblage that, if undisturbed, may lead
to further differentiation, should these variants be able to persist for
the necessary millenniums in our rapidly changing world. Adjacent
to the Fringillidae I place the other groups that obviously are closely
allied to them. Attempts to arrange the avian families with the Cor-
vidae and their allies in the terminal position, as accepted in various
earlier classifications, and as followed now by Mayr and Amadon,
because of supposed more advanced development of the brain ap-
pear to me quite uncertain, particularly in view of our decidedly
limited information in this field. Should this idea be coupled with be-
lief in superior mental reactions in the corvine assemblage, I would
consider this more an anthropomorphic interpretation than one sup-
ported by scientific fact.

The formation of the group names has been one of recent interesting
discussion. The suffixes -idae and -inae for families and subfamilies
are accepted rather universally so that they do not require examina-
tion. In view of the limited number of species covered in ornithology
I see no point in the introduction of tribes as another category between
the subfamily and the genus. This may be useful to entomologists
with their tens of thousands of species, but seems unnecessary and

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE 13

cumbersome with birds. In some of the more comprehensive avian
genera there are groups of species more closely allied to one another
than to their fellows, but the taxonomist may discuss these at need as
groups without imposing another burden on a classification that now
is highly divided. For the group names above the family level, I be-
lieve it preferable to use suffixes that allow immediate identification of
the rank, coupled with a stem that, like the family name, is based on a
current generic term. Where ordinal and subordinal names are both
formed as Latin plurals there is possibility of confusion.

The detailed classification that follows shows as its main improve-
ments our better understanding of some of the peculiar birds found
in Australia and New Zealand, and some additional historical data of
importance in our slowly growing knowledge of extinct forms in the
fossil record. Stresemann, and more recently Amadon and Mayr,
have made many valuable suggestions as to the relationships of nu-
merous peculiar genera that in part are still of uncertain position. It
remains intriguing to observe the vast amount of information still
to be secured before our basis for final classification may be considered
complete.

October I, 1951.

Class Aves, Birds.
Subclass Archaeornithes, Ancestral Birds.
Order Archaeopterygiformes, Archaeopteryx, Archaeornis.
Family Archaeopterygidae, Archaeopteryx (fossil),
Archaeornithidae, Archaeornis (fossil).
Subclass Neornithes, True Birds.
Superorder Odontognathae, New World Toothed Birds.
Order Hesperornithiformes, Hesperornithes.
Family Hesperornithidae, Hesperornis, Hargeria (fos-
sil).
Enaliornithidae,** Enaliornis (fossil).
Baptornithidae, Baptornis (fossil).
Order Ichthyornithiformes, Ichthyornithes.
Family Ichthyornithidae, Ichthyornis (fossil).
Apatornithidae, Apatornis (fossil).

84 Position provisional. Lambrecht, Handbuch der Palaeornithologie, 1933,
Pp. 255-261, unites the Baptornithidae with this group, which he considers closely
related to the loons and grebes.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Superorder Impennes, Penguins.

Order

Sphenisciformes, Penguins.
Family Spheniscidae, Penguins.

Superorder Neognathae, Typical Birds.

Order

Caenagnathiformes, Caenagnathus.
Family Caenagnathidae, Caenagnathus *° (fossil).
Struthioniformes, Ostriches.
Family Eleutherornithidae, Eleutherornis (fossil).
Struthionidae, Ostriches.
Rheiformes, Rheas.
Family Rheidae, Rheas.
Casuariiformes, Cassowaries, Emus.
Family Casuariidae, Cassowaries.
Dromiceidae, Emus.
Dromornithidae, Dromornis (fossil).
Aepyornithiformes, Elephantbirds.
Family Aepyornithidae, Aepyornis (fossil and extinct).
Dinornithiformes, Moas.
Family Dinornithidae, Moas (fossil and extinct).
Anomalopterygidae, Anomalopteryx, Emeus,
and Allies (fossil and extinct).
Apterygiformes, Kiwis.
Family Apterygidae, Kiwis.
Tinamiformes, Tinamous.
Family Tinamidae, Tinamous.
Gaviiformes, Loons.
Family Gaviidae, Loons.
Colymbiformes, Grebes.
Family Colymbidae, Grebes.
Procellariiformes, Albatrosses, Shearwaters, Petrels,
and Allies.
Family Diomedeidae, Albatrosses.
Procellariidae, Shearwaters, Fulmars.
Hydrobatidae, Storm Petrels.
Pelecanoididae, Diving Petrels.
Pelecaniformes, Tropic-birds, Pelicans, Frigate-birds,
and Allies.

Suborder Phaéthontes, Tropic-birds.

Family Phaethontidae, Tropic-birds.

85 This interesting genus, listed tentatively in the above superorder, possibly

is not avian.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—WETMORE I5

Suborder Pelecani, Pelicans, Boobies, Cormorants, Snake-
birds.
Superfamily Pelecanoidea, Pelicans and Allies.
Family Pelecanidae, Pelicans.
Cyphornithidae, Cyphornis, Palaeochenoides
(fossil).
Superfamily Suloidea, Boobies, Cormorants, and Allies.
Family Pelagornithidae, Pelagornis (fossil).
Sulidae, Boobies, Gannets.
Elopterygidae, Elopteryx, Eostega, Actiornis
(fossil).
Phalacrocoracidae, Cormorants.
Anhingidae, Snake-birds
Suborder Fregatae, Frigate-birds.
Family Fregatidae, Frigate-birds.
Suborder Odontopteryges, Odontopteryx, Pseudodontornis,
Cladornithes (fossil).
Family Odontopterygidae, Odontopteryx (fossil).
Pseudodontornithidae,*® Pseudodontornis (fos-
sil).
Cladornithidae, Cladornis, Cruschedula (fossil).
Order Ciconiiformes, Herons, Storks, and Allies.
Suborder Ardeae, Herons, Bitterns.
Family Ardeidae, Herons, Bitterns.
Cochleariidae, Boat-billed Herons.
Suborder Balaenicipites, Whale-headed Storks.
Family Balaenicipitidae, Whale-headed Storks.
Suborder Ciconiae, Storks, Ibises, Spoonbills.
Superfamily Scopoidea, Hammerheads.
Family Scopidae, Hammerheads.
Superfamily Ciconioidea, Storks.
Family Ciconiidae, Storks, Jabirus.
Superfamily Threskiornithoidea, Ibises.
Family Threskiornithidae, Ibises, Spoonbills.
Suborder Phoenicopteri, Flamingos.
Family Agnopteridae, Agnopterus (fossil).
Scaniornithidae, Scaniornis, Parascaniornis
(fossil).
Phoenicopteridae, Flamingos.

86 Position not certain; see Lambrecht, Handbuch der Palaeornithologie, 1933,
PP. 305-308.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.) 117,

Order Anseriformes, Screamers, Ducks, Geese, Swans.
Suborder Anhimae, Screamers.
Family Anhimidae, Screamers.
Suborder Anseres, Ducks, Geese, Swans.
Family Paranyrocidae, Paranyroca (fossil).
Anatidae, Ducks, Geese, Swans.
Order Falconiformes, Vultures, Hawks, Falcons.
Suborder Cathartae, New World Vultures.
Superfamily Neocathartoidea, Neocathartes.
Family Neocathartidae, Neocathartes (fossil).
Superfamily Cathartoidea, New World Vultures.
Family Cathartidae, New World Vultures.
Teratornithidae, Teratornis, Cathartornis (fos-
sil).
Suborder Falcones, Secretarybirds, Hawks, Falcons.
Superfamily Sagittarioidea, Secretarybirds.
Family Sagittariidae, Secretarybirds.
Superfamily Falconoidea, Hawks, Falcons, and Allies.
Family Accipitridae, Hawks, Old World Vultures,
Harriers.
Pandionidae, Ospreys.
Falconidae, Falcons, Caracaras.
Order Galliformes, Megapodes, Curassows, Pheasants, Hoat-
zins.
Suborder Galli, Megapodes, Curassows, Grouse, Pheasants.
Superfamily Cracoidea, Megapodes, Curassows.
Family Megapodiidae, Megapodes.
Gallinuloididae, Gallinuloides (fossil).
Cracidae, Curassows, Guans, Chachalacas.
Superfamily Phasianoidea, Grouse, Pheasants, Turkeys.
Family Tetraonidae, Grouse.
Phasianidae, Quails, Pheasants, Peacocks.
Numididae, Guineafowl.
Meleagrididae, Turkeys.
Suborder Opisthocomi, Hoatzins.
Family Opisthocomidae, Hoatzins.
Order Gruiformes, Cranes, Rails, and Allies.
Suborder Mesoenatides, Roatelos, Monias.
Family Mesoenatidae, Roatelos, Monias.
Suborder Turnices, Bustard-quails, Hemipodes.
Family Turnicidae, Bustard-quails.
Pedionomidae, Plain-wanderers.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE 17

Suborder Grues, Cranes, Limpkins, Trumpeters, Rails.
Superfamily Gruoidea, Cranes, Limpkins, Trumpeters.
Family Geranoididae, Geranoides (fossil).
Eogruidae, Eogrus (fossil).
Gruidae, Cranes.
Aramidae, Limpkins.
Psophiidae, Trumpeters.
Superfamily Ralloidea, Rails.
Family Orthocnemidae,** Orthocnemus, Elaphrocne-
mus (fossil).
Rallidae, Rails, Coots, Gallinules.
Suborder Heliornithes, Sun-grebes.
Family Heliornithidae, Sun-grebes.
Suborder Rhynocheti, Kagus.
Family Rhynochetidae, Kagus.
Suborder Eurypygae, Sun-bitterns.
Family Eurypygidae, Sun-bitterns.
Suborder Phororhaci, Phororhacos and Allies.
Family Phororhacidae, Phororhacos and Allies (fos-
sil).
Psilopteridae, Psilopterus and Allies (fossil).
Brontornithidae, Brontornis, Liornis, and Allies
(fossil).
Opisthodactylidae, Opisthodactylus (fossil).
Cunampatidae, Cunampaia (fossil).
Suborder Cariamae, Cariamas and Allies.
Family Bathornithidae, Bathornis (fossil).
Hermosiornithidae, Hermosiornis, Procariama
(fossil).
Cariamidae, Cariamas.
Suborder Otides, Bustards.
Family Otididae, Bustards.
Order Diatrymiformes, Diatryma, Omorhamphus, and Allies.
Family Diatrymidae, Diatryma (fossil).
Gastornithidae, Gastornis, Remiornis (fossil).
Order Charadriiformes, Shorebirds, Gulls, Auks.
Suborder Charadrii, Shorebirds.
Superfamily Jacanoidea, Jacanas.
Family Jacanidae, Jacanas.

87 Position provisional. See Lambrecht, Handbuch der Palaeornithologie, 1933,
PP. 490-493.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 17

Superfamily Charadrioidea, Plovers, Sandpipers, and Al-
lies.

Family Rhegminornithidae, Rhegminornis (fossil).
Rostratulidae, Painted Snipe.
Haematopodidae, Oystercatchers.
Charadriidae, Plovers, Turnstones, Surfbirds.
Scolopacidae, Snipe, Woodcock, Sandpipers.
Recurvirostridae, Avocets, Stilts.
Presbyornithidae, Presbyornis (fossil).
Phalaropodidae, Phalaropes.

Superfamily Dromadoidea, Crab-plovers.

Family Dromadidae, Crab-plovers.

Superfamily Burhinoidea, Thick-knees.

Family Burhinidae, Thick-knees.

Superfamily Glareoloidea, Pratincoles, Coursers.

Family Glareolidae, Pratincoles, Coursers.

Superfamily Thinocoroidea, Seed-snipe.
Family Thinocoridae, Seed-snipe.
Superfamily Chionidoidea, Sheath-bills.
Family Chionididae, Sheath-bills.

Suborder Lari, Gulls, Terns, Skimmers.

Family Stercorariidae, Skuas, Jaegers.
Laridae, Gulls, Terns.
Rynchopidae, Skimmers.

Suborder Alcae, Auks.
Family Alcidae, Auks, Auklets, Murres.
Order Columbiformes, Sand-grouse, Pigeons, Doves.
Suborder Pterocletes, Sand-grouse.
Family Pteroclidae, Sand-grouse.
Suborder Columbae, Pigeons and Doves.

Family Raphidae, Dodos, Solitaires.
Columbidae, Pigeons, Doves.

Order Psittaciformes, Lories, Parrots, Macaws.

Family Psittacidae, Lories, Parrots, Macaws.

Order Cuculiformes, Plantain-eaters, Cuckoos.
Suborder Musophagi, Plantain-eaters.
Family Musophagidae, Plantain-eaters.
Suborder Cuculi, Cuckoos, Roadrunners, Anis.
Family Cuculidae, Cuckoos, Roadrunners, Anis.

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE 19

Order Strigiformes, Owls.
Family Protostrigidae, Protostrix (fossil).
Tytonidae, Barn Owls.
Strigidae, Owls.
Order Caprimulgiformes, Oilbirds, Goatsuckers.
Suborder Steatornithes, Oilbirds.
Family Steatornithidae, Oilbirds.
Suborder Caprimulgi, Frogmouths, Goatsuckers.

Family Podargidae, Frogmouths.

Nyctibiidae, Potoos.
Aegothelidae, Owlet-frogmouths.
Caprimulgidae, Goatsuckers.
Order Apodiformes, Swifts, Hummingbirds.
Suborder Apodi, Swifts.

Family Aegialornithidae,** Aegialornis (fossil).

Apodidae, Swifts.
Hemiprocnidae, Crested Swifts.
Suborder Trochili, Hummingbirds.
Family Trochilidae, Hummingbirds.
Order Coliiformes, Collies.
Family Coliidae, Colies.
Order Trogoniformes, Trogons.
Family Trogonidae, Trogons.
Order Coraciiformes, Kingfishers, Bee-eaters, Rollers, Horn-
bills.
Suborder Alcedines, Kingfishers, Todies, Motmots.
Superfamily Alcedinoidea, Kingfishers.
Family Alcedinidae, Kingfishers.
Superfamily Todoidea, Todies.
Family Todidae, Todies.
Superfamily Momotoidea, Motmots.
Family Momotidae, Motmots.
Suborder Meropes, Bee-eaters.
Family Meropidae, Bee-eaters.
Suborder Coracii, Rollers, Hoopoes.

Family Coraciidae, Rollers.
Brachypteraciidae, Ground-rollers.
Leptosomatidae, Cuckoo-rollers.
Upupidae, Hoopoes.
Phoeniculidae, Wood-hoopoes.

38 Placed here provisionally. See Lambrecht, Handbuch der Palaeornithologie,
1933, pp. 623-624.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Suborder Bucerotes, Hornbills.
Family Bucerotidae, Hornbills.
Order Piciformes, Jacamars, Barbets, Toucans, Woodpeckers.
Suborder Galbulae, Jacamars, Barbets, Toucans.
Superfamily Galbuloidea, Jacamars, Puffbirds.
Family Galbulidae, Jacamars.
Bucconidae, Puffbirds.
Superfamily Capitonoidea, Barbets, Honey-guides.
Family Capitonidae, Barbets.
Indicatoridae, Honey-guides.
Superfamily Ramphastoidea, Toucans.
Family Ramphastidae, Toucans.
Suborder Pici, Woodpeckers.
Family Picidae, Woodpeckers, Piculets.
Order Passeriformes, Perching Birds.
Suborder Eurylaimi, Broadbills.
Family Eurylaimidae, Broadbills.
Suborder Tyranni, Ovenbirds, Tyrant Flycatchers, and Al-
lies.
Superfamily Furnarioidea, Ovenbirds, Woodhewers, and
Allies.
Family Dendrocolaptidae, Woodhewers.
Furnariidae, Ovenbirds.
Formicariidae, Ant-thrushes.
Conopophagidae, Ant-pipits.
Rhinocryptidae, Tapaculos.
Superfamily Tyrannoidea, Tyrant Flycatchers, Pittas, and
Allies.
Family Cotingidae, Cotingas.
Pipridae, Manakins.
Tyrannidae, Tyrant Flycatchers.
Oxyruncidae, Sharpbills.
Phytotomidae, Plant-cutters.
Pittidae, Pittas.
Acanthisittidae, New Zealand Wrens.
Philepittidae, Asities, False Sunbirds.
Suborder Menurae, Lyrebirds.
Family Menuridae, Lyrebirds.
Atrichornithidae, Scrub-birds.
Suborder Passeres, Songbirds,
Family Alaudidae, Larks.
Palaeospizidae, Palaeospiza (fossil).

NO. 4 REVISED CLASSIFICATION FOR BIRDS OF WORLD—-WETMORE 2I

Hirundinidae, Swallows.

Campephagidae, Cuckoo-shrikes.

Dicruridae, Drongos.

Oriolidae, Old World Orioles.

Corvidae, Crows, Magpies, Jays.

Cracticidae, Bell Magpies, Australian Butcher-
birds.

Grallinidae, Magpie-larks.

Ptilonorhynchidae, Bowerbirds.

Paradiseidae, Birds of Paradise.

Paradoxornithidae, Parrotbills, Suthoras.

Paridae, Titmice.

Sittidae, Nuthatches.

Hyposittidae, Coral-billed Nuthatches.

Certhiidae, Creepers.

Chamaeidae, Wren-tits.

Timaliidae, Babbling Thrushes.

Pycnonotidae, Bulbuls.

Cinclidae, Dippers.

Troglodytidae, Wrens.

Mimidae, Thrashers, Mockingbirds.

Turdidae, Thrushes.

Zeledoniidae,*® Wren-thrushes.

Sylviidae, Old World Warblers.

Regulidae, Kinglets.

Muscicapidae, Old World Flycatchers.

Prunellidae, Accentors, Hedge-sparrows.

Motacillidae, Wagtails, Pipits.

Bombycillidae, Waxwings.

Ptilogonatidae, Silky Flycatchers.

Dulidae, Palm-chats.

Artamidae, Wood-swallows.

Vangidae, Vanga Shrikes.

Laniidae, Shrikes.

Prionopidae, Wood-shrikes.

Cyclarhidae, Pepper-shrikes.

Vireolaniidae, Shrike-vireos.

Callaeidae, Wattled Crows, Huias, Saddlebacks.

Sturnidae, Starlings.

39 For notes on Zeledonia see Ridgway, U. S. Nat. Mus. Bull. 50, pt. 4, 1907,

p. 885.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Meliphagidae, Honey-eaters.

Nectariniidae, Sunbirds.

Dicaeidae, Flower-peckers.

Zosteropidae, White-eyes.

Vireonidae, Vireos.

Coerebidae, Honey-creepers.
Drepanididae, Hawaiian Honey-creepers.
Parulidae, Wood Warblers.

Ploceidae, Weaver-finches.

Icteridae, Blackbirds, Troupials.
Tersinidae, Swallow-tanagers.
Thraupidae, Tanagers.
Catamblyrhynchidae, Plush-capped Finches.
Fringillidae, Grosbeaks, Finches, Buntings.

}

'SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 5

" ANNOTATED LIST OF BIRDS OF BARRO
a * ~ COLORADO ISLAND, PANAMA
| CANAL ZONE

BY,
EUGENE EISENMANN

sens

( Pusticatios 4038)

Tah CITY OF WASHINGTON - |
ae "PUBLISHED BY THE SMITHSONIAN INSTITUTION
ie _ FEBRUARY ve 1952, bi

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 5

PNINOTATED LIST OF BIRDS OF BARRO
COLORADO ISLAND, PANAMA
CANAL ZONE

BY.
EUGENE EISENMANN

(PUBLICATION 4058)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
FEBRUARY 7, 1952

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ANNOTATED LIST: OF BIRDS’ OFS BARRO
COLOKADO ISLAND, PANAMA’ CANAL ZONE

By EUGENE EISENMANN

Few localities in tropical America have been visited by so many
naturalists as the biological station on Barro Colorado Island. Yet
after nearly 30 years since the island was reserved for scientific
purposes, even the inventory of its resident bird life is probably
incomplete. A number of species collected in nearby parts of the
Canal Zone still remain unrecorded, and almost every year students
report one or two birds previously unknown from the island. This
will surprise no one having field experience with the type of lofty
tropical forest covering the island, where some woodland species are
represented by few individuals, and others, though more numerous,
are likely to be overlooked unless one is familiar with their notes.

In regard to the behavior and ecological relations—or, indeed, such
fundamentals as breeding habits and season—of even most of the
common species little is known and less has been published. Our
ignorance applies to such species generally, not merely to their status
on Barro Colorado Island. Thus the facilities of this very accessible
area afford students extraordinary opportunities for adding to our
knowledge of tropical birds.

LOCATION AND ADMINISTRATION

Barro Colorado is the largest island in Gattn Lake, Panama Canal
Zone, lying opposite the station of Frijoles on the Panama Railroad.
Though situated on the Atlantic slope of the Continental Divide and
less than 15 miles from the Caribbean Sea, it is only 20 miles from
the Pacific. It was formed by the damming of the Chagres River
for the Canal construction which converted the lowlands into Gatun
Lake and the hilltops into islands. The ship channel through the
Lake, in view of the laboratory, affords a fascinating contrast be-
tween the crisp modernity of ocean-going steamers and the wild,
green islands and shores, against an ever-changing background of
distant mountains, blue sky, and billowing clouds.

1Frijoles station, where one gets the launch for the island, is only a short
train ride from the cities of Coldn and Panama. At present, daily airplanes

from Miami reach Panama in 4} hours.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 5

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 1L7

Barro Colorado rises to a height of about 452 feet above the lake
surface and 537 feet above sea level. Irregular in outline, with over
25 miles of shore line formed by many deeply indented bays and
inlets, the island has a maximum diameter of about 3 miles and an
area of some 3,609 acres, or 5.6 square miles. But its surface habita-
ble for animal life is much greater than if it were level ground, for
the whole is broken by numerous drainage ravines whose slopes are
covered with luxuriant vegetation. Despite the well-kept trails, cross-
ing the island on foot takes far more time and effort than the air-line
diameter might suggest.

Barro Colorado Island, under its official title of Canal Zone Bi-
ological Area, is at present administered by the Smithsonian Institu-
tion. The island was set aside as a biological station on April 17, 1923,
through the efforts of Thomas Barbour and of James Zetek, who
from the start has served as resident manager. It was operated by
the Institute for Research in Tropical America until 1940. Then, on
July 2, 1940, Congress gave the area a permanent status, but World
War II suspended normal activities. It was placed under the
Smithsonian Institution by Presidential order on July 16, 1946.
Since then visitors and students have again been welcomed and pro-
vided the means of becoming acquainted with the biological wealth
of a humid tropical forest under comfortable and healthful conditions.

The station provides laboratory and library facilities, as well as
the basic needs of food and lodging, in an environment of unusual
beauty and interest. Well-marked trails crisscross the forest, and
at various points on the periphery small houses are strategically
located, so that the night may be spent in the more remote portions
of the island.

Those wishing to visit Barro Colorado Island should communicate
with the Secretary of the Smithsonian Institution, Washington 25,
D. C., or with the Resident Manager, Canal Zone Biological Area,
Drawer C, Balboa, C. Z.

ECOLOGICAL CONDITIONS

Barro Colorado Island is in the humid division of the Lower Tropi-
cal Zone, but there is a distinct dry season. The average annual
rainfall (1925-1949) is about 107 inches, almost all of it between
May and December ; November is the wettest month. The four dry-
season months, January, February, March, and April, all together
supply on the average only about 7 inches (Zetek, 1950). The rainy
season usually begins late in April and ends late in December. Even

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 3

in this season precipitation comes chiefly at night and in the after-
noon. During the dry season, while rainfall may be less than 1 inch
in some months, the high relative humidity (about 75 percent) suffices
to keep the forest green, though in February and March some of
the higher trees drop their leaves and produce masses of brilliant
flowers. Daily temperatures range between 70° and go° (average
below 80°) ; variation is between night and day rather than between
seasons (Zetek, 1948). During the dry season strong northeast trade
winds blow; during the rainy season there is generally a pleasant
breeze from the lake.

Except for a few acres of clearing, the island is entirely wooded.
About half is mature forest; the remainder is in various stages of
succession (Kenoyer, 1929).

For its size and relatively uniform environment Barro Colorado
Island has a strikingly rich and varied flora and fauna. Standley
(1933) lists 1,259 species of plants. Of vertebrates, other than
birds, 56 species of mammals, 62 reptiles, 33 amphibians, and 22 fishes
are reported (Zetek, 1951). The regularly resident larger mammals
include monkeys (4 species), puma, ocelot, tayra, coati, kinkajou,
deer (2 species), peccary (2 species), tapir, sloths (2 species), ant-
eaters (2 species), and agouti (Enders, 1935).

In all, 306 species of birds are here listed as satisfactorily identified
from the island ; 52 are migrants from North America and I is proba-
bly a migrant from South America. The remainder are “resident”
in the sense that they breed within the Republic of Panama, though
some of them, including certain water birds that one sees daily, defi-
nitely do not nest on Barro Colorado Island, and others merely appear
occasionally from the mainland. Although direct breeding evidence
(in the form of nests or fledglings) is presently available for fewer
than I00 species, it is probable, on the basis of their year-round
presence and known habits, that about 200 species breed more or
less regularly on the island.2, The population is by no means static.
This can readily be noted around the clearing: from day to day one
observes different species, and birds breeding there one year may be
wholly absent the next at the very same period. Diurnal movements

2The number of species one may expect to see on a short visit is smaller,
as conditions in the forest make many birds very elusive. For example, during
a 2-week period in June and July (when there are no migrants), covering the
laboratory clearing and perhaps a third of the trails, I ordinarily note between
120 and 125 species; over a hundred in or about the clearing. In fact, without
leaving the clearing, on almost any day I can count (including birds heard)
55 to 65 species; but it takes a good deal of effort to raise a day’s list beyond 75
at this season.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

to and from the island, by such birds as the parrots, are very obvious.
To what extent there are also seasonal movements by the “resident”
birds is not known, but that they occur in some species seems likely.

Though essentially a humid neotropical forest environment, from
the bird student’s viewpoint the island habitats may conveniently be
considered under four headings.

1. The clearing.—As here used this includes a narrow strip extend-
ing from the dock or entrance cove up the hillside to and around the
laboratory buildings, and connecting with another narrow strip (some-
times called the plantation) running some 300 yards along the lake.
This is largely a grassy area, interspersed with small trees (chiefly
cultivated citrus) and banana plants. Crowding closely around the
clearing a lofty forest rises sharply. About the clearing may be
observed the greatest variety of birds. In fact, from there most of
the species recorded on the island have at one time or another been
seen or heard. A number of species requiring relatively open country,
otherwise absent from the island, have become established or occa-
sionally appear. The densely grown borders are favored by thicket
inhabitants. The extensive “edge effect’’ exerts its well-known attrac-
tiveness. Best of all, the hillside location enables one to view the lake,
the sky, the grassy slope, and the upper levels of the nearby forest.
Many birds of the forest canopy are rarely noted except when flying
across the clearing or moving in the adjacent treetops. Here, unlike
the forest, bird activity is apparent throughout the day. Much smaller
cleared areas are maintained at the summit of the island (where there
is an observation tower) and at a few scattered points along the lake
shore, but these are relatively insignificant, though they have attracted
some birds absent from the forest.

2. The mature or primary forest covers about half the island, par-
ticularly the western portion. Though some exists near the laboratory,
it is most impressive in the more level southwestern quarter. This
forest is characterized by three arboreal strata, the upper canopy
being between 75 and 100 feet, with scattered overtopping trees rising
125 feet or more (Allee, 1926b). There is relatively little undergrowth.
Where best developed, the mature forest, with its dim filtered light
and columnar structure, produces a cathedral solemnity. The con-
stant hum of the cicadas soon fades into an unnoticed background,
and the effect on a windless day is of silence—a silence interrupted
at long intervals by the screeches of parrots or toucans, the roaring
or chattering of monkeys, or the rapid passage of a mixed band of
birds, perhaps accompanying a train of army ants. The mature forest
seems to be preferred by most of the larger birds and mammals. The
flowering and fruiting trees attract many smaller species, but it is

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 5

usually impossible to see, and often even to hear, such birds in the
canopy. Except early in the morning, and to a less extent late in the
afternoon, few birds are noticeable in the forest.

3. The secondary forest occupies the remainder of the island. It
varies from areas that 25 years ago were clearings to forest well
advanced toward maturity. The trees are lower and the canopy less
closed than in the mature forest, so more light reaches the floor. As
a result the undergrowth is thicker—in some places impenetrable
without cutting. Epiphytes grow nearer the ground; ropelike, woody
lianas and smaller vines are abundant. The secondary forest has a
much wilder appearance than the mature forest, and more closely
approximates the popular conception of a tropical jungle. Some
birds are found only in the lighter woodland, others favor the denser
tangles. But the correlation of neotropical birdlife to the forest suc-
cession remains largely an untilled field.

4. The water borders—While these habitats vary, depending on
exposure, maturity of the forest, and other factors, they have in
common the favorable combination of standing water, abundant light,
and large trees. The light encourages the growth of a green wall of
vegetation down to the water’s edge, providing haunts for thicket
dwellers. In addition to birds for which the proximity of water is
a requirement, this environment attracts a number of other species,
particularly flycatchers, usually considered clearing birds. The most
beautiful water borders are the margins of the esteros, narrow inlets
(actually drowned stream valleys), deeply indenting the island, and
best explored by cayuco, the native dug-out canoe. Along the quiet
esteros aquatic plants grow luxuriantly, and the “edge effect’ is
apparent in the variety of birds. Extensive marshy vegetation is
found only on the southwest margin of the island, where, protected
from the waves raised by the trade winds and the wake of steamers,
little grassy islets have formed around projecting tree stumps, thus
affording homes for gallinules, rails, and jacanas.

Gatun Lake itself, despite its richly forested shores, seems strangely
unattractive to waterfowl. While masked ducks breed in its narrower
reaches, none has ever been reported from the vicinity of Barro
Colorado. Pied-billed grebes and anhingas occur, but the only swim-
ming bird commonly seen is the ubiquitous cormorant.

ORNITHOLOGICAL WORK RELATING TO BARRO COLORADO
ISLAND

In the first few years of the station’s existence, careful collecting,
chiefly by Dr. Frank M. Chapman and Dr. Josselyn Van Tyne, deter-
mined the basic avifauna. For many years now collecting of birds

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

has been prohibited, except in very special cases. Most additions to
the bird list since 1928 have been based on sight observations.

In 1929 Dr. Chapman, in his first delightful book on Barro Colorado,
“My Tropical Air Castle’—which every prospective visitor should
read—listed some 200 species of birds personally recorded by him
in the course of four dry seasons (late December to early April).
In his equally enjoyable sequel, “Life in an Air Castle” (1938),
published after 12 such seasons on the island, he appended a list of
251 species, including records by other observers. Dr. Chapman also
wrote elaborate studies on the breeding habits of the manakin, Mana-
cus vitellinus (1935), and of the oropéndola, Zarhynchus wagleri
(1928).

It would serve no purpose to list all the bird students who have
briefly visited Barro Colorado Island. Many of them are mentioned
below among those who have contributed to the preparation of this
list. The station has served to introduce students from all over the
world to neotropical wildlife. Aside from Dr. Chapman, those orni-
thologists who have spent most time there are Dr. Alfred O. Gross,
Dr. Alexander F. Skutch, and Dr. Josselyn Van Tyne. Dr. Gross
has written several articles describing the nesting of various Barro
Colorado birds (1927, 1929, 1930, 1950). Dr. Van Tyne published
a monograph on the toucan, Ramphastos brevicarinatus (1929), and
shorter accounts of breeding by other species (1926, 1929, 1950).
Dr. Skutch, the most assiduous student of Middle American bird
behavior, has published many papers containing life-history material
obtained on Barro Colorado Island (1931-1951). Much of the Barro
Colorado breeding data here given is derived from his observations.

Partly on the basis of studies on Barro Colorado, Mrs. Bertha B.
Sturgis wrote the “Field Book of Birds of the Panama Canal Zone”
(1928), which gives descriptions, with many illustrations, of most
species known to occur in the Canal Zone. It is, I believe, the only
pocket-sized descriptive bird book in English covering any part of
the continental neotropics.

My own interest in Panama bird life dates back to my childhood.
I first visited Barro Colorado in 1937. Since then I have sojourned
there for periods between June and September in 1938, 1945, 1948,
1949, 1950, and 1951. Though I have had dry-season experience in
Panama, all my stays on the island have been during the rainy season ;
thus my personal observations are somewhat complementary to the
dry-season studies of Dr. Chapman.

In an effort to make the list as complete as present information per-
mits, an appeal for Barro Colorado Island data was made through the

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISEN MANN 7

columns of the Auk, Condor, Wilson Bulletin, and Linnaean News-
Letter, as well as by direct correspondence. The following supplied me
with helpful information: Dr. A. A. Allen, Robert S. Arbib, Jr., Dr. F.
Bourliere, Geoffrey Carleton, Dr. Nicholas E. Collias, G. W. Cottrell,
Jr., Dr. David E. Davis, Miss Hazel Ellis, Mrs. Gladys Gordon Fry
(including observations of her companions, Mrs. A. Edey, Mrs. M.
Edey, Mrs. L. J. Francke, and Mrs. French), Thomas Gilliard, Dr.
A. O. Gross, Dr. David Harrower, Mrs. Dorothy M. Hobson (includ-
ing data of her companion Miss C. A. Moore), Thomas Imhof, Dr.
Lawrence Kilham, Robert Laughlin, Dr. Frederick W. Loetscher, Jr.,
Philip Longenecker, Drs. Lorus J. and Margery Milne, Dr. Charles
W. Quaintance, Dr. Robert T. Scholes, Dr. Alexander F. Skutch, Dr.
Ken W. Stott, Jr., Dr. Josselyn Van Tyne, Jay A. Weber, and Dr.
Alexander Wetmore.

Both published and unpublished data, including that contained in
the species index on Barro Colorado Island, have been critically
examined, bearing in mind the fact that several of the observers had
had no prior experience with neotropical birds. Some doubtful reports
have been omitted, in almost all cases with the approval of my cor-
respondent.* I have examined specimens in the collections of the
American Museum of Natural History and on Barro Colorado Island,
and Dr. Van Tyne and Dr. Gross kindly checked for me the species
represented in their respective collections.

No one who has enjoyed the facilities of Barro Colorado can fail
to acknowledge his gratitude to James Zetek, the resourceful and
hospitable guardian of the island, but for whom there would be no
biological station. In connection with the preparation of this paper
I wish particularly to note the help of Dr. Gross, who originally urged
me to this task, of Dr. Van Tyne, who generously provided me with
much valuable data and criticism, and of Dr. Wetmore, who as Secre-
tary of the Smithsonian Institution has had the island under his sym-
pathetic supervision; all three read the manuscript and gave me
excellent suggestions.

THE, ANNOTATED. LIST

In this list species are marked with an asterisk when actual speci-
mens taken on Barro Colorado Island are known. In view of the policy
against collecting on the island, the absence of such specimens has little

3In a few instances the footnotes refer to certain reports by competent ob-
servers of species whose reported presence, while unlikely or subject to possible
misidentification, seems worth mentioning. :

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

significance ; actually of the 306 species here listed all (except Phaeo-
progne tapera) have been collected within nearby parts of Panama,
and almost all within the Canal Zone.

Technical names.—The nomenclature followed is basically that of
Griscom’s Panama checklist (1935), with modifications suggested by
more recent studies of Hellmayr, Friedmann, Peters, Wetmore, and
Zimmer.

English names.—These are intended to be comprehensive names
applicable to the species as a whole—not merely to the Canal Zone
subspecies—and thus sometimes differ from names that have been used
for the local race.‘

Unfortunately no published list of English species names exists for
Middle American birds, as the older authorities generally provided
only subspecies names, which usually did not indicate conspecific rela-
tionship.» Until some committee acts, the individual writer or student
is forced to make his own selection.

For migrants from the area of the A. O. U. Check-list of North
American Birds the species names here used are those so far adopted
by the A. O. U. Committee.

For tropical species the literature was examined to find a name
appropriate to each species (not misleading as to any of its races or
confusing with the name of other species), and, if possible, with some
associative or recognition value. Hence names suggestive of some
character of the appearance, behavior, habitat, or general distribution
of the species were favored, and patronymics and lesser geographic
designations were avoided (see Eisenmann and Poor, 1946). Prefer-
ence was given to appropriate names used for the species or one of
its subspecies in Ridgway’s Birds of North and Middle America or
in Hellmayr’s Catalogue of Birds of the Americas ; and when a choice
existed among the names of several subspecies, that of the nominate

4 To facilitate reference, where a name selected is materially different from
that used in the Sturgis Field Book (1928), that name is indicated in paren-
theses.

5 This practice (often criticized) not only obscured relationships but use-
lessly multiplied the number of disparate names to be memorized. It has been
particularly burdensome in the neotropics, where most species are divided into
many local and intergrading subspecies, so that there is frequently uncertainty
as to the subspecific status of the local population. The amateur, for whom Eng-
lish names are primarily intended, often has no name available unless he pre-
tends to make a subspecific discrimination that cannot possibly be made in the
field. The use of appropriate comprehensive names avoids these difficulties,
facilitates recognition throughout the range, and can aid in identification.

;
|
|
:
|

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 9

subspecies was favored. Resort was also had to the writings of others.®
Effort was made to avoid novelty, although some names were modified
to prevent confusion or misleading effect.

Status.—Status and habitat on Barro Colorado Island are indicated
where known, but our information is still very tentative and incom-
plete. In the case of migrants, the spotty Barro Colorado data have
been supplemented with migration dates from Panama, derived from
published records, from labels on specimens in the American Museum,
and from sight records of my own or of a few other observers.

Notes.—The descriptions of calls and song are not intended as a
complete account of the notes of any species. They are given primarily
as an aid to identification, in the full realization that verbal interpreta-
tions tend to be extremely personal. In a few instances I include
interpretations by other writers.

Breeding.—All breeding dates from Barro Colorado reported to me
have been summarized. Since this information is meager, I have added
supplemental data (also very incomplete) of nesting dates elsewhere
in the Canal Zone and nearby parts of Panama. On the information
presently available it can be said that, while there are birds nesting on
Barro Colorado at all times of the year, the greatest number seem to
breed between March and June. The breeding periods of Central
American birds are still inadequately known, but helpful accounts
have been prepared by Harrower (1936a) and Skutch (19502).

Descriptions.—Certain species recorded from Barro Colorado
Island are not described in the Sturgis Field Book (1928). For the
convenience of visitors I have supplied a brief diagnosis of such birds,
other than migrants from North America.

Order TINAMIFORMES: Tinamous
Family TINAMIDAE: Tinamous

*Tinamus major castaneiceps: Great Tinamou (Chestnut-headed).

Common in the forest; heard daily from the clearing at dusk and
dawn, occasionally at other times. Notes: A sad, flutelike whistle of
two long-drawn, tremulous notes, the second beginning slightly higher
but sliding down; sometimes the first note is repeated once or twice
before the second is given; sometimes the 2-noted phrase is twice
uttered. Breeding: February 26, 1934, 8 young just able to fly

6 Particularly Sutton’s Mexican Birds (1051), the recent Distributional
Check-list of the Birds of Mexico (Friedmann et al., 1950), and publications of
Beebe, Gould, and Skutch.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 117

(Davis) ; February 28, 1938, 2 eggs (Gilliard) ; April 28, 1926, 6 eggs
(Van Tyne); June 21, 1951, 4 eggs (Milne, Eisenmann) ; June 26,
1927, 4 eggs (Van Tyne) ; June 30, 1948, 2 downy young (Eisen-
mann).

*Crypturellus soui panamensis: Little Tinamou (Pileated).

Status uncertain ; formerly not uncommon near the laboratory ; not
seen in recent years, though reported calling, by Wetmore, on April
24, 1947, April 5, 1948. Prefers second growth and forest edge.
Notes: A series of tremulous, ascending whistles, increasing in volume
and rapidity, then usually dying away suddenly with a few falling
notes. Most often heard early in the morning or late in the afternoon.
Breeding: March 16, 1926, 2 eggs (Van Tyne) ; July 13, 1927, 2 eggs,
hatched July 29, 1927 (Gross and Van Tyne) ; August 4, 1925, 2 eggs
(Gross).

Order COLYMBIFORMES: Grebes
Family COLYMBIDAE: Grebes

*Podilymbus podiceps antarcticus: Pied-billed Grebe.

Male in breeding condition collected August 5, 1927 (Van Tyne) ;
others seen March 8, April 14, and June 3, 1927 (Van Tyne).

Order PELECANIFORMES: Totipalmate Swimmers
Family PELECANIDAE: Pelicans

*Pelecanus occidentalis carolinensis: Brown Pelican.
Small groups fly daily over the lake on their way across the isthmus.
They nest on islands in the Gulf of Panama.®

Family PHALACROCORACIDAE: Cormorants

*Phalacrocorax olivaceus olivaceus: Olivaceous Cormorant (Bra-
zilian).
Very common on the lake, especially on projecting tree trunks.

7 The least grebe (Colymbus dominicus) occurs, and probably breeds, on
Gattin Lake, but has never been reported from Barro Colorado Island.

8 Four blue-faced boobies (Sula dactylatra) are reported by Mrs. G. G.
Fry flying together over the Lake on February 14, 1940, “within easy identifi-
cation distance.” As no storm had occurred, this is very surprising, for the
species, though often noted off Col6n Harbor, is distinctly pelagic.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—-EISENMANN Tr

Family ANHINGIDAE: Anhingas

* Anhinga anhinga leucogaster: Anhinga.
Not common; occasionally seen in the laboratory cove and the nar-
row esteros.
Family FREGATIDAE: Frigatebirds

Fregata magnificens rothschildi: Magnificent Frigatebird.

Small groups fly daily over the lake on their way across the isthmus.
They nest on islands in the Gulf of Panama.

Order CICONIIFORMES: Herons, Storks, Ibises, and Allies
Family ARDEIDAE: Herons

Ardea herodias: Great Blue Heron.

Individuals occur at all seasons, including May, June, July, and
August (Van Tyne, Eisenmann, Loetscher, Longenecker, Stott).
Wetmore believes that some of these are migrant individuals of the
northern great blue heron and that some may be of another subspecies.

Casmerodius albus egretta: Common Egret (American).

Casual on the island. Seen March 11, 1946, February 22, 1948,
April 5, 1948, February 28, 1951 (Wetmore).

*Hydranassa tricolor ruficollis: Tricolored Heron (Louisiana).

Individuals infrequently noted (Chapman). Female collected Octo-
ber 30, 1927 (Gross).

*Florida caerulea: Little Blue Heron.

Individuals seen throughout the year along the lake shore ; the heron
most frequently noted in the laboratory cove ; most birds are immature.

Leucophoyx thula thula: Snowy Egret.

February 27, 1940 (Mrs. A. Edey and Mrs. M. Edey). The rarity
here of this common Canal Zone species is probably caused by the
absence of mud flats.

*Butorides virescens: Green Heron.

Birds of the genus Butorides are the most numerous of the family
on the island. Almost all students have reported seeing green herons.
Wetmore writes: “Two forms of this species undoubtedly occur, one

12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

resident and one migrant, but field records are confused owing to the
fact that the two differ only in size. The northern virescens winters
in the Canal Zone (specimens identified from near Gattn and Taber-
nilla) ; the smaller race maculatus, presumably resident, is also found
(specimens seen from near Gattn). Birds, supposedly maculatus,
have been reported breeding on the Island by Chapman and others:
March 24, 3 eggs (Gilliard) ; April 28, 1935, 2 eggs (Skutch). Addi-
tional breeding specimens are needed to determine the true composition
of the resident population.”

*Butorides striatus patens: Striated Heron.

Common along the wooded lake margins.® Van Tyne (1950) attrib-
utes to this form all breeding birds of this genus collected by him on
the island. Breeding: April 11, 1927, male taken in full breeding con-
dition; July 28, 1925, nest, 2 eggs; August I1, 1925, 2 half-grown
young (Van Tyne).

Agamia agami: Agami or Chestnut-bellied Heron.

Estero west of Salud Point, May 10, 1935 (Skutch). A fairly
large (length, 32 inches) slender heron. Adult is bottle green above
with lavender crown and back plumes, and largely maroon and chest-
nut below. Immature is dark brown above with slaty crown, tail, and
wing tips, and has creamy-buff underparts with dusky streaks on the
sides.

Nycticorax nycticorax hoactli: Black-crowned Night Heron.

Occasional (Chapman). Breeds in Canal Zone, March 21 (Hal-
linan).

Tigrisoma mexicana: Bare-throated Tiger-heron (Cabanis’s).

“Observed rarely. No specimens” (Chapman).*°

9 Some individuals approach B. virescens in being quite brownish-necked. As
to this Wetmore says: “Adults are never deep chestnut on the neck like B.
virescens, but show various shadings of lighter brown toward the purely gray-
necked B. s. striatus, which is found in eastern Panama. Immature birds may
be separated in the field with difficulty from B. v. maculatus.”

10 The common tiger-heron (T. 1. lineatum), which has a feathered throat
and the adults of which have the head and neck chestnut, may be expected to
occur.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 13

Ixobrychus exilis: Least Bittern.

Infrequently seen in marshy islets on southwestern shore (Chap-
man). The northern exilis may occur in winter ; the South American
erythromelas may breed ; both have been collected in the Canal Zone.

Family CICONIIDAE: Storks

Mycteria americana: Wood Ibis.

One seen flying over the island on February 12, 1940 (Mrs. Fry,
Mrs. Francke, and Mrs. Teague).*?

Order FALCONIFORMES: Birds of Prey
Family CATHARTIDAE: Vultures

*Sarcoramphus papa: King Vulture.

Not common, but seen from time to time, often flying over the clear-
ing. Rarely more than one individual, but four soaring together Feb-
ruary 8, 1950 (Wetmore), and September 18, 1951 (Bourliere).

*Coragyps atratus: Black Vulture.
One or two individuals generally can be seen daily.

*Cathartes aura aura: Turkey Vulture.

Four to eight individuals generally can be seen daily. Large flocks
of migrating birds, apparently one of the northern forms, have been
noted: February 25-April 5 (Chapman, Cottrell).

Family ACCIPITRIDAE: Hawks

*Elanoides forficatus yetapa: Swallow-tailed Kite.

Not uncommon ; from time to time small groups are seen flying over
the clearing. A flock of 20 noted June 30, 1949 (Eisenmann).

*Leptodon cayanensis: Gray-headed Kite.

Adult female collected, perched on a forest trail, August 18, 1927
(Van Tyne). Fairly large (length, 18-24 inches), bill very hooked.

11 Fight all-dark unidentified ibises, apparently of the family Threskiornithidae,
were seen flying over by Mrs. Fry February 9, 1940. While the green ibis
(Mesembrinibis cayennensis), a bottle-green bird with greenish bill and legs,
is the only all-dark ibis so far taken in the Canal Zone, two or three other
such species may possibly occur.

T4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL Lv

Adults: Above blackish with a contrasting slaty-gray head ; underparts
white, sometimes tinged with pearl gray, tail with two or three whitish
bands. Immatures: Similar, but head mainly white with black crown,
a creamy white collar, and white underparts. There are also melanistic
phases with blackish head and streaking below.

*Chondrohierax uncinatus uncinatus: Hook-billed Kite (Red-collared
Hawk).
Occasionally noted near the edge of the clearing. Collected May
18, 1926 (Van Tyne) ; seen July 10, 11, 1950 (Eisenmann).

*Harpagus bidentatus fasciatus: Double-toothed Kite.

Occasionally seen at the edge of the clearing and in the forest.
Rather small (length, 14 inches). Adults: Above (including sides of
head) slaty; below largely chestnut, breast and abdomen usually bar-
red with white, the bars often edged with gray. Immatures: Above
brown; below yellowish white, throat and chest sparsely streaked
with dusky, belly and flanks barred with brown. Notes: A high shrill
pee-yip, also pseeyp, pseeyp. Breeding: Nest high in Bombacopsis
tree at clearing edge, June 29, 1951; egg taken by toucan July 4
(Laughlin, Eisenmann).

*Tctinia plumbea: Plumbeous Kite.

Uncommon. Collected April g (male, breeding condition), April 22
(female), 1926 (Van Tyne). Seen April 25, 1926 (pair), three flying
over the forest August 17, 1927 (Van Tyne); February 25, 1950
(Kilham) ; March 19, 1950 (Cottrell).

*Buteo platypterus platypterus: Broad-winged Hawk.

Winter visitant. A flight of over 250 hawks, some definitely this
species,?? noted March 21, 1933 (Carleton). Late date April 6, 1950
(Wetmore).

Buteo magnirostris petulans: Roadside or Insect Hawk (Large-billed).

Occasional around the clearing; the common open-country hawk
over much of Panama. Collected at Frijoles (Gilliard).

*Leucopternis albicollis costaricensis: White Hawk (Ghiesbrecht’s).

One or two birds generally frequent the forest edge near the labora-
tory, and often perch in the open, permitting close approach.

12 Swainson’s hawk (B. swainsoni) migrates in large flocks over the Canal
Zone, but has not yet been reported from the island.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 15

Leucopternis semiplumbea: Semiplumbeous Hawk.

“Not common” (Chapman). Rather small (length, 15 inches).
Adults: Above lead gray ; below white, usually with a few fine throat
streaks; wings black, tail black with one white band. Immatures:
Similar, but more streaked below and tail with two white bands.

Hypomorphnus urubitinga ridgwayi: Greater Black Hawk, or Urubi-
tinga.

One seen March 8, 1949 (Wetmore). Resembles the next species
but is larger and heavier (length, 26 inches) with decidedly longer
legs. Adults: Differ also in having pure white upper tail coverts, thighs
distinctly barred with white, and slaty preocular facial skin contrast-
ing with yellow cere. Immatures: Also like those of next species, but
head more buffy with less blackish streaking and throat generally
unstreaked buffy.

Buteogallus anthracinus: Common Black Hawk or Crab Hawk.
Adult seen April 6, 1948 (Wetmore). Commonest in the coastal
areas of Panama.

*Morphnus guianensis: Common Crested Eagle.

Collected March 30, 1936 (R. J. Niedrach and A. C. Rogers). Very
large (length, 32-37 inches), with pointed crest. Adults: Above slaty
with lighter tipping to crest ; below chiefly white, more or less barred
on breast and belly with rufous or brown, chest gray to brown.
Immatures: Above ashy white with brown marbling, head (except
dusky tips to crest) and entire underparts white ; tail dusky with irreg-
ular lighter bands. Melanistic examples also occur.

*Morphnus taentatus: Banded Crested Eagle.

One seen in the forest February 8, 1950 (Wetmore). This rare bird
may be a color phase of the preceding species (Lehmann, 1943; Hell-
mayr and Conover, 1949). It differs in having a blackish chest, with
the breast and belly broadly barred black and white.

Harpia harpyja: Harpy Eagle.
Seen 1924 (Barbour), January 8, 1947 (Quaintance), June 22, 1950
(Stott). ‘

Spizastur melanoleucus: Black-and-white Eagle-hawk.

“Rare” (Chapman). Large (length, 21-25 inches), crested, tarsus
feathered to the toes. Adults: Head, neck, and underparts white, crest

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

and nape with a few black markings; rest of upper parts blackish,
wings and tail barred with paler. Immatures: Similar, but dark areas
more brownish and wing coverts narrowly tipped with whitish.

*Spizaétus ornatus vicarius: Ornate or Crested Eagle-hawk.

Collected August 17, 1927 (Van Tyne) ; seen soaring over an open-
ing on Pearson trail February 8, 1950 (Wetmore). Large (length,
24-49 inches), conspicuously crested. Adults: Above blackish; ear
coverts and sides of neck and chest tawny rufous; a broad black
mustache mark below eye; rest of underparts white, belly barred
with black. Jmmatures: Above browner, entire head and neck buffy
white, rest of underparts white, with a few black spots and black
barring on sides.

Spizaétus tyrannus serus: Black or Tyrant Eagle-hawk.

Seen perched in forest at edge of clearing June 28, 1949 (Eisen-
mann, Gross). Large (length, 26-30 inches), conspicuously crested.
Adults: Black, with some white on crown; white barring on thighs,
under wings coverts, under tail coverts, and tail. Immatures: Above
dark brown, nape and crest white with black tips, wings barred and
spotted with black and white; below largely blackish, with white on
center of throat and chest and on flanks, chest streaked with brown.

Circus cyaneus hudsonius: Marsh Hawk.
Regular winter visitant (Chapman).

Geranospiza nigra nigra: Blackish Crane-hawk or Frog-hawk.

Several sight records around the clearing: February 13, 1940 (Mrs.
A. Edey, Mrs. M. Edey, Mrs. H. Scherman), January 8, 1945 (Allen),

July 19, 1945, July 2, 1949 (Eisenmann).

Family PANDIONIDAE: Ospreys

Pandion haliaéius carolinensis: Osprey.
Regular winter visitant. Though not a breeder, individuals are seen
in Panama throughout the year.

Family FALCONIDAE: Falcons
*Micrastur ruficollis interstes: Barred Forest-falcon (Cassin’s Bar-
red Hawk).

Adult female collected May 1, 1926, in forest on Barbour-Lathrop
trail (Van Tyne). ‘Not uncommon” (Chapman). A forest species.

———— -—-—-

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISEN MANN iy

*Micrastur semitorquatus naso: Collared or Pied Forest-falcon
(Black-and-white Hawk).

Collected February 2, 1926 (Chapman and Potter). Occasionally
seen in the forest below the canopy. March 11, 1946 (Wetmore).
Notes: A series of deliberate laughing notes, hah, hah, hah, or a lower
cuh, cuh, cuh. Breeding: July 20, 1949, two fully grown young sitting
in a tree (Loetscher).

*Daptrius americanus guatemalensis: Red-throated Caracara.

Fairly common in the forest, wandering about in small bands, which
will approach an intruder with noisy protests. Notes: A loud, raucous
ca-ca-ca, cd-o, sometimes cd-ak, ca-cd-o.

Falco peregrinus anatum: Peregrine Falcon (Duck Hawk).

Uncommon migrant and winter visitant: February 12, 1937 (Gil-
liard) ; November 8, 1944 (Allen).

*Falco albigularis albigularis: Bat Falcon.

A pair or two probably reside on the island, for one or two birds
can often be seen flying over the clearing or along the lake shore.
Notes: kee-kee-kee-kee. Breeding: In Panama near the Canal Zone
a pair seen feeding a full-grown juvenal, June 26, 1949 (Eisenmann).

Order GALLIFORMES: Gallinaceous Birds
Family CRACIDAE: Curassows and Guans

*Crax rubra rubra: Great Curassow (Great Panama).

Recorded only by Van Tyne: collected male (of two seen) on April
18, 1927; male seen June 25, 1925, and on three occasions in the
spring of 1926. All were found “in the big forest, one not far behind
the Laboratory, the others up towards the center of the island.”

*Penelope purpurascens aequatorialis: Crested Guan.

Common in the forest, occasional in the clearing. Usually found
in small bands in the trees, calling loudly. Notes: A somewhat varia-
ble metallic yelping, quenk, quenk, quenk, or keelp, keelp, keelp. In
January before sunrise drumming is reported by Chapman.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

*Ortalis garrula cinereiceps: Chestnut-winged Chachalaca (Gray-
headed).

Fairly common in the lighter growth, especially near the lake shore ;
occasional around the clearing. Notes: A loud, repeated chack, chack,
chack.

Family PERDICIDAE: Partridges and Quails

Odontophorus gujanensis marmoratus: Marbled Wood-quail.

Formerly occurred in undergrowth at edge of the clearing; not
reported in recent years. Notes: A repeated, very rapid, ringing,
musical call syllabized as corcorovado, corcorovado, corcorovado
(Chapman) or burst the bubble, burst the bubble, burst the bubble
(Skutch).

Order GRUIFORMES: Cranes, Rails, and Allies
Family RALLIDAE: Rails

*Aranudes cajanea cajanea: Gray-necked Wood-rail (Cayenne).

Not common; frequents damp woodland. Notes: Pairs duet, usu-
ally,at night “resembling an aged couple singing in shaky, quavering
voices” (Chapman), a “loud series of short gurgling laughs” (AI-
drich), or killi, killi, killi followed by cow, cow, cow (Young).

*Laterallus albigularis albigularis: White-throated Crake.

“Common in grassy shores and islands” of the southwestern part
(Chapman). Notes: “A short but loud, groaning snore” (Chapman).

*Porphyrula martinica: Purple Gallinule.

Pairs well distributed where small marshy areas occur along the
bays and estero margins. Notes: A variety of clucking, cackling, and
guttural sounds, as well as a snapping of the bill (Gross and Van
Tyne, 1929). Breeding: March 14, 1929, 2 young about a week old
(Chapman) ; May 15, 1935, nest, 5 eggs near Salud Point (Skutch) ;
July 13, 1925, nest, 4 eggs near laboratory cove, hatched August 2-3
(Gross and Van Tyne); November 5, 1927, young in natal down
near Frijoles, and many juvenals about 3 months old seen or collected
October and early November 1927 (Gross and Van Tyne).

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 19

Family HELIORNITHIDAE: Sun-grebes

*Heliornis fulica: Sun-grebe (American Finfoot).

Local and uncommon; all reports are from the coves and esteros
of Gigante Bay on the south side of the island. Collected August 18,
1927 (Van Tyne) and November 6, 1927 (Gross). Seen March 15,
1933 (Carleton); January 3, 1937, cove near Drayton house
(Skutch) ; March 9, 1949, cove at end of Shannon trail (Wetmore) ;
July 8, 1950, cove near Drayton house (Eisenmann, Dr. and Mrs.
F. Ryan).

Family EURYPYGIDAE: Sun-bitterns

*Eurypyga helias major: Sun-bittern.

Rare; collected March 15, 1926, in a creek bed deep in the forest
on Shannon trail (Van Tyne); “‘seen on the lake and along small
forest streams tributary to it” (Sturgis).

Order CHARADRIIFORMES: Shorebirds, Gulls, Auks
Family JACANIDAE: Jacanas

*Jacana jacana hypomelaena: Wattled Jacana (Black).

Fairly common in aquatic vegetation, especially on the southwest
margin of the island. Breeding: March 24, 1937, four young about a
day old (Gilliard).

Family SCOLOPACIDAE: Snipes and Sandpipers

*Actitis macularia: Spotted Sandpiper.

Regular migrant and winter visitant. Late date: May 12, 1926
(collected Van Tyne). There are Canal Zone records for every
month except June, and in Panama from July 25 to May 23.

Family LARIDAE: Gulls and Terns

Larus atricilla: Laughing Gull.

Common migrant and winter visitant on the lake; occasional at
other times. Collected on May 17, 1926, at Frijoles (Van Tyne).
Several seen June 22, 27 and July 3, 1951 (Eisenmann). Many imma-
ture birds can regularly be found in Panama waters throughout the
year.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

Larus pipixcan: Franklin’s Gull.

One seen July 3, 1951, with the preceding species, on the lake
between Barro Colorado Island and Frijoles. Primarily a migrant
in Panama, but immatures sometimes linger over the summer, as
noted repeatedly in June-July 1951.

Sterna hirundo: Common Tern.

Seen July 1, 1950, and June 25 and July 3, 1951 (Eisenmann).
Individuals, apparently immature, can regularly be found in Panama
waters even in June and July. Jewel collected a male from a flock
and a lone immature female on Gatttn Lake on December 3, 1911,
and June 9, 1912, respectively (Stone, 1918).

*Thalasseus maximus maximus: Royal Tern.

Regular migrant and winter visitant over the lake. Collected May
28, 1927, seen July 19 and 23, 1927, August 16, 1927 (Van Tyne),
and June 25, 1951 (Eisenmann). Many immature birds regularly
occur in Panama waters throughout the year.

Thalasseus sandvicensis acuflavidus: Sandwich Tern (Cabot’s).
November 7, 1944, seen over lake (Allen).

*Chlidonias niger surinamensis: Black Tern.

Regular migrant, occasional in summer: collected November 12,
1927 (Gross). Seen November 29, 1945 (R. C. Murphy), June 28,
1949, six birds (Eisenmann, Gross), July 10, 13, 1949 (Loetscher).
Considerable numbers in nonbreeding plumage remain in Panama
Bay during the summer.

Order COLUMBIFORMES: Pigeonlike Birds

Family COLUMBIDAE: Pigeons

*Columba speciosa: Scaled Pigeon.

“Not uncommon” (Chapman). Collected by Van Tyne. An
arboreal species of lighter forest and edge.

*Columba cayennensis pallidicrissa: Pale-vented Pigeon.

Fairly common arboreal species of the lighter woodland, especially
along the lake shore. Notes: A mournful kiik-tu-codoo.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 21

*Columba nigrirostris: Short-billed Pigeon.

Fairly common arboreal forest species. Notes: A “melodious ‘Je
t’adore’’”’ (Chapman) ; to my ear, ho, cu-cu-codoo.

Columbigallina minuta elaeodes: Plain-breasted Ground-dove.

March 24, 1950, one seen in the clearing (Cottrell). An open-
country bird.

Columbigallina talpacoti rufipennis: Ruddy Ground-dove.

Occasionally occurs in the clearing (Weber, Skutch). Common in
the more open parts of the Canal Zone and Panama.

*Claravis pretiosa pretiosa: Blue Ground-dove.

A pair may usually be found at the edge of the clearing, and
occasionally elsewhere in the lighter woods. Primarily a thicket bird,
both terrestrial and arboreal. Breeding: Nests with eggs found in
the Canal Zone in August (Arbib), and female about to lay in April
(Stone, 1918).

Leptotila verreauxi verreauxi: White-tipped Dove (Verreaux’s).

A pair or two can be found in or around the clearing, and occasion-
ally in other fairly open areas. Favors thickets and small trees in
open country, but feeds chiefly on the ground. Notes: A soft, inquir-
ing hod-oo or hoo-hoo, hoo-oo. Breeding: March 1947 (Quaintance) ;
June 30, 1950, incubating (Eisenmann) ; July 1949 (Gross) ; August
5, 1942, two eggs (Mrs. Hobson) ; nests with eggs September 18,
1925, October 8, 1927 (Gross).

*Leptotila cassinui cassinu: Gray-chested Dove (Cassin’s).

Well distributed throughout the forest, occasionally seen at the
edge of the clearing. Frequents the ground and lower growth. The
gray head contrasting with the brown nape and back help to distin-
guish this species from the preceding. Breeding: February-April
1935 (Skutch) ; April 12, 1937 (Gilliard) ; July 30, 1934 (Arbib).
In the Canal Zone also May and September (Stone, 1918).

Geotrygon montana: Ruddy Quail-dove.

Not uncommon on the ground in the forest. Notes: “A soft deep
coo” (Skutch) ; to my ear a low humming mmmm, repeated after
a half-second pause.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 7

Order PSITTACIFORMES: Parrots
Family PSITTACIDAE: Parrots

*Brotogeris jugularis jugularis: Orange-chinned or Tovi Parakeet.

Common; gathering in substantial flocks in the clearing after the
breeding season, particularly late in the afternoon. More than 100
birds gathered in one tree every evening in September 1938, then
flew off to the mainland to roost for the night. Notes: A high screech-
ing chatter. Breeding: January-May (Van Tyne).

*Pionus menstruus: Blue-headed Parrot.

Common in the forest; seen daily in the clearing, usually flying
over, but sometimes perching. Distinguished in flight from the Ama-
gona parrots by higher-pitched calls and deeper, freer wing strokes ;
the short wing strokes of the Amazona group suggest the flight of
ducks. Notes: A harsh, high keéweenk, keéweenk, keéweenk. Breed-
ing: April 12, 1937, young in nest (Gilliard) ; April 6, 1950, in stub
in the water (Wetmore).

Amazona autumnalis salvini: Red-lored Parrot (Salvin’s).

Common in the forest; seen daily flying over the clearing, particu-
larly morning and afternoon. Notes: Harsh screeches syllabized as
chikak, chikék, oordk, oordk, ooérk (Chapman).

Amazona ochrocephala panamensis: Yellow-headed Parrot (Panama).

Rare or casual; a bird of the lighter Pacific woodlands. Reported
by Sturgis; also July 16, 1945, 2 perched at the edge of the clearing
(Eisenmann) ; March 17, 1950, 3 or 4 on Barbour trail (Cottrell).

*Amazona farinosa inornata: Mealy Parrot (Plain-colored).

Common in the forest; pairs seen daily flying over the clearing.
Notes: Louder, but less raucous than those of the preceding, rendered
as stop it, stop it, quick quick quick and yelp, yelp, yelp, cha-cha-cha
(Chapman).

Order CUCULIFORMES: Cuckoolike Birds
Family CUCULIDAE: Cuckoos

Coccyzus americanus: Yellow-billed Cuckoo.

Migrant seen April 30, 1951 (Collias). Recorded in Panama
September 30—November 19.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISEN MANN 23

*Piaya cayana thermophila: Squirrel Cuckoo.

Individuals are well distributed where there is forest edge or high
thickets. Notes: A dry chick-kaw; a loud trrt-trrt-irrt-irrt; a ringing,
almost whistled whep, repeated five to eight times ; a strongly accented
kyékerah or wipperah; also piscdtaqua piscdtaqua (Chapman) ; also
a sharp peek and a lower chirr (Mrs. Hobson). Breeding: Nest in
mango tree beside the kitchen, May 1935 (Skutch).

Crotophaga major: Greater Ani.

Local; small groups are spottily distributed in shrubbery along
the lake edge of various esteros. In July 1950 I noted three groups
on the south shore in different esteros of Gigante Bay and three on
the north shore at coves between Fuertes Estero and Salud Point.
Notes: Most characteristic is a bubbling prrrr or brrrr, resembling
the sound produced by expelling air forcibly from between tightly
closed lips; also a lighter whirrrr; and a very different, almost mam-
mal-like growl grrrwa or grrrr.

*Crotophaga anit: Smooth-billed Ani.

A small group, which has on occasion included as many as 11 birds,
can usually be found in the lower part of the clearing or on Slothia
Island ; rarely elsewhere along the shore; 2 seen northwest of Armour
trail-end, June 25, 1951. Notes: Most characteristic is a whining
oooo-eeeek or oooo-eeeelk. Breeding: January and February (Chap-
man, Gilliard) ; February-May 1935, repeated nestings, all destroyed
(Skutch) ; June 1929 (Skutch).

Tapera naevia excellens: Striped Cuckoo.

Occasional in the clearing ; a bird of open-country thickets. Notes:
Melancholy, long-drawn whistles. One phrase is of two notes, the
second note a half tone higher ; another phrase is of five or six notes,
the next to the last highest and longest. Breeding: Said to be parasitic.

*Dromococcyx phasianellus rufigularis: Pheasant Cuckoo.

Rare ; collected March 13, 1926 (Van Tyne) ; seen (Chapman). A
bird of thick undergrowth.

*Neomorphus geoffroyi salvini: Rufous-vented Ground-cuckoo (Sal-
vin’s).
Collected by Chapman and Potter January 7, 1926; seen at the
beginning of Barbour-Lathrop trail, March 22, 1935 (Skutch). A
terrestrial bird of forest undergrowth.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS. LE7

Order STRIGIFORMES: Owls
Family STRIGIDAE: Owls **

Otus choliba luctisonus: Choliba Screech Owl.

Occurs about the clearing; a bird of open woods, clearings, and
gardens. “Not common” (Chapman) ; April 23, 1947 (Wetmore) ;
February 1950 (Kilham). Notes: A short series of quick purring or
bubbling notes ending in a higher catlike whine.**

Lophostrix cristata stricklandi: Crested Owl (Strickland’s).

Two sight reports: Armour trail, January 18, 1929 (Robert S.
Woods) ; near laboratory, July 10, 1948 (Longenecker).

*Pulsatrix perspicillata chapman: Spectacled Owl.

Seen from time to time near the laboratory. Notes: A. A. Allen
attributes to this species a series of low hoots, of almost machine-gun
rapidity, of which he made a sound recording. Chapman (1929)
thought that a very different, deep, deliberate woof-woof-woof, often
heard before dawn, might be uttered by this species. Neither Allen
nor Chapman saw the bird when it was calling.

*Ciccaba virgata virgata: Mottled Wood-Owl (Cassin’s Barred Owl).
“One specimen; status unknown” (Chapman). A forest owl.

Order CAPRIMULGIFORMES: Goatsuckers and Allies
Family CAPRIMULGIDAE: Goatsuckers

*Chordeiles acutipennis micromeris: Lesser Nighthawk (Texas).

Immature female collected October 30, 1927 (Gross), one of a
number of migrants, large flocks having been noted from late Sep-
tember on (Gross, 1930a). This specimen, now in the American Mu-
seum of Natural History (wing, 173 mm., tail, 95.3 mm.), is iden-
tified by Wetmore as micromeris, the Central American race. Other
races, including terensis of southwestern United States, must also
occur in migration. On September 14, 1938, I saw a flock of night-
hawks, but as the birds were not calling and were flying high, and

18 The status of the island owls is very uncertain owing to lack of knowl-
edge of their voices.

14 The vermiculated screech owl (Oitus guatemalae vermiculatus), which dif-
fers in lacking both the distinct dark facial rim and the herringbone pattern
below, may also occur, for it has been collected at Frijoles.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 25

the common nighthawk (C. minor) also migrates over the Canal
Zone, the species could not be determined.

*Nyctidromus albicollis intercedens: Pauraque.

Often heard and occasionally seen around the clearing. Notes:
A hoarse whistled cor-weéee-oo or por-weéeecer; also a bup-bup-bup
or hip-hip-hip on a single tone. Breeding: March 1935 in laboratory
clearing, April 30, 1935, on Slothia Island (Skutch).

*Caprimulgus carolinensis: Chuck-will’s-widow.

Migrant ; one found dead November 1931 on steps leading to labora-
tory (in Barro Colorado Island collection).

Family NYCTIBIIDAE: Potoos

Nyctibius grandis: Great Potoo.
“Rare” (Chapman). A little-known nocturnal woodland species.
Notes: Reported as oorrr and oorrroo in Surinam (Haverschmidt,

1948).

Nyctibius griseus panamensis: Common Potoo (Panama).

“Not uncommon” (Chapman). A nocturnal woodland species.
Notes: A melancholy series of loud, rich, long-drawn wailing notes
(usually six), going down scale (Chapman).

Order APODIFORMES: Swifts and Hummingbirds
Family APODIDAE:* Swifts

Panyptila cayennensis: Swallow-tailed Swift.

Occasional over the clearing: September 12-16, 1938, 2 to 5 birds
daily (Eisenmann) ; April 6, 1949 (Wetmore) ; July 23, 1950 (Eisen-
mann); June 23, 1951 (Scholes, Eisenmann). Breeding: Found
nesting in the Canal Zone in July on Ancon Hospital piazza and Pedro
Miguel locks (Greenway, 1934). A velvety-black, fork-tailed swift,
with white throat, nape, and sides of rump.

15 Swifts of the genus Chaetuwra are often seen over the clearing, but no
specimens have been taken. Chapman (1938) apparently assumed them to be
C. chapmani. As five species of Chaetura have been collected in or near the
Canal Zone (Rogers, 1939) and their separation in the field is very difficult,
sight identifications seem unreliable—at least until more is known regarding
their local status.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Family TROCHILIDAE: Hummingbirds

Glaucis hirsuta affinis: Rufous-breasted Hermit (Lesser Hairy).

Not uncommon, especially in wild plantain (Heliconia) thickets
and around banana plants. Breeding: In the Canal Zone nests found
in April and May (Stone, 1918).

Threnetes ruckeri ventosus: Band-tailed Barbed-throat (Rucker’s
Hermit).

Occasionally noted around Heliconia and banana plants.

Phaethornis guy coruscus: Green Hermit (Bangs’s).
Not uncommon, in damp woods and Helicoma thickets.

*Phaethornis superciliosus 1® cassinii: Long-tailed Hermit (Nica-
raguan).

Fairly common; in damp woods and Heliconia thickets. Breeding:
Nests at tip of palm leaf; August 5, 1925, 2 eggs (Gross), June 24,
IQ51, 2 eggs, 1 hatched June 26 (Milne, Eisenmann). In the Canal
Zone, nest with eggs in May (Stone, 1918).

*Phaethornis longuemareus saturatus: Little Hermit (Dusky).

Occasional in the clearing, not uncommon in the forest ; always but
a few feet above the ground.

Florisuga mellivora mellivora: White-necked Jacobin (Jacobin Hum-
mingbird).
Common about the flowering shrubs of the clearing. Breeding:
January 10, 1937, nest with eggs (Gilliard).

Anthracothorax nigricollis nigricollis: Black-throated Mango.

Not uncommon around the clearing. Breeding: December, January,
February, and March (Chapman, Skutch, Allen).

Lophornis delatirei lessoni: Rufous-crested Coquette (De Lattre’s).

Females seen near the laboratory June 30, 1948, and June 28—July
5, 1949 (Eisenmann).

16 Zimmer (1950) considers P. malaris, not P. superciliosus, to be the nomi-
nate race of the wide-ranging species which includes the Middle American
forms.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 27

Chlorostilbon canivetu assimilis: Fork-tailed Emerald (Allied).

“Not common” (Chapman). A bird partial to gardens, clearings,
and the more open country of the Pacific slope. Breeding: In the
Canal Zone, nests found in January and November (Sturgis).

*Thalurania furcata venusta: Common Wood-nymph (Colombian).
Common around the clearing and in the forest. Breeding: March
2, March 25, 1935, nests in the forest (Skutch).

*Damophila julie panamensis: Violet-bellied Hummingbird (Panama).

One of the most common hummingbirds around the flowering
shrubs of the clearing, at least during June-September. Chapman
regarded it as “not common” during the dry season.

Lepidopyga coeruleogularis coeruleogularis: Sapphire-throated Hum-
mingbird (Duchassain’s).
Occasional around the laboratory. A common species in more open
country.

Hylocharts eliciae: Blue-throated Goldentail (Elicia’s).

Males seen at the laboratory July 5, 1948, July 2, 1949, June-July
1950 (Eisenmann).

*Amazilia amabilis costaricensis: Blue-chested Hummingbird
(Lovely).
Male collected March 12, 1926 (Van Tyne).

Amazilia edward edward: White-bellied Hummingbird (Wilson’s).

Males seen at the laboratory July, 2, 6, 1948, and July 4, 6, 1950
(Eisenmann). Generally a bird of more open country.

*Amazilia tzacatl tzacatl: Rufous-tailed Hummingbird (Rieffer’s).

The most common hummingbird around the clearing. Breeding:
December 1930 (Skutch) ; July 10, 1950, collecting nesting material
(Eisenmann). In the Canal Zone, nests found in April and May
(Stone, 1918).

*Chalybura buffonii micans: White-vented Plumeleteer (Buffon’s).
Occasional around the laboratory.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, It7

*Heliothryx barroti: Purple-capped Fairy (Barrot’s).
Common in flowering trees around the clearing. Breeding: April
1924, nest attached to side of vertical hanging liana (Sturgis).

Heliomaster longirostris longirostris: Long-billed Starthroat.
Male seen January 27, 1951, near the laboratory (Collias). Pri-
marily an open-country species.

Order TROGONIFORMES: Trogons
Family TROGONIDAE: Trogons

*Trogon massena hoffmanni: Slaty-tailed Trogon (Massena).

Common in the forest, often seen in the clearing. Notes: A loud
cah, cah, cah, interminably repeated at the rate of about two a second.
Breeding: In old termite nests March-June (Chapman, Gilliard) ;
July 13, 1927, nest with young (Gross).

Trogon melanurus macroura: Black-tailed Trogon (Large-tailed).
Very few records. “Rather rare” (Chapman) ; woods near labora-
tory March, April, 1935 (Skutch).

*Trogon rufus tenellus: Black-throated Trogon (Graceful).
Common in the forest; often seen at the edge of the clearing.
Notes: cow, repeated three or four times, with a pause between notes,
which come at the rate of about one a second; also a sharp, repeated
chirr. Breeding: In low decaying stumps; April 1935, 2 nests
(Skutch) ; April 20, 1951, 2 eggs (Collias) ; June 18, 1951, 2 eggs
(Milne), 1 young had hatched June 21 (Milne, Eisenmann).

Trogon violaceus concinnus: Violaceous Trogon (Gartered).

Common in the lighter forest ; often seen at the edge of the clear-
ing. Notes: A rather soft cow, repeated at a steady pace of about two
a second, 10 to 15 times. Breeding: February 25, 1950, 2 eggs
in cavity near top of 6-foot stump (Kilham) ; April 1936, building in
termite nest (Chapman) ; June-July 1950, building in termite nest
(Eisenmann).

*Trogon viridis chionurus: White-tailed Trogon.
Common in the lighter forest; often seen in the clearing. Notes:
A rather soft coo, repeated at first slowly but accelerating into a roll,

NO. § BIRDS OF BARRO COLORADO ISLAND—EISENMANN 29

which sometimes ends with a few slower notes; occasionally a bird
will vary the usual call with a simple kuh, kuh, kuh, kuh. Breeding:
March II, 1947, in termite nest (Wetmore) ; April, building in ter-
mite nest (Chapman) ; June 27, 1950, feeding young in termite nest
(Stott, Eisenmann).

Order CORACIIFORMES: Kingfishers, Motmots, and Allies
Family ALCEDINIDAE: Kingfishers

*Megaceryle torquata torquata: Ringed Kingfisher.
Common along the lake shore.

Megaceryle alcyon alcyon: Belted Kingfisher.
Common winter visitant.

*Chloroceryle americana isthmica: Green Kingfisher.
Most common kingfisher. Notes: A clicking trit-trit-trit.

Chloroceryle nda: Green-and-rufous Kingfisher.

Seen April 17, 1942, between Frijoles and Barro Colorado Island
(Imhof).

*Chloroceryle amagona mexicana: Amazon Kingfisher.
Fairly common along the lake shore.

Chloroceryle aenea aenea: Pygmy Kingfisher (Least Green).

Occasionally seen in the narrower bays and inlets: near dock
July 21, 1945 (Eisenmann) ; estero near Drayton house August 22,
1948 (Longenecker) ; at mouth of stream east of Armour trail-end,
April 6, 1951 (Collias).

Family MOMOTIDAE: Motmots

*Electron platyrhynchum minor: Broad-billed Motmot.

Fairly common in the forest; often heard from the laboratory in
the early morning. Notes: A single loud and resonant, but somewhat
nasal cwa or cah, repeated with pauses of 2 to 3 seconds; occasionally
run together as cah-cah-cah.

*Baryphthengus ruficapillus semirufus: Rufous Motmot.

Common in the forest; heard several times daily from the labora-
tory, particularly in the early morning. Notes: The basic phrase is a

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

mellow, rhythmic hod-too-too or hé-hoo-hoo, with the accent on the
first syllable, often repeated, and echoed by other individuals. A
common call sounds like hoérro. The hooting may be varied in pitch
or time, and is frequently accelerated into a roll.

Momotus momota conexus: Blue-crowned Motmot (Small-billed).

“Occasional ; irregular” (Chapman). A bird of more open wood-
land, collected at Frijoles (Gilliard).

Family BUCCONIDAE: Puffbirds

*Notharcus macrorhynchus hyperrhynchus: White-necked Puffbird
(Dyson’s).
Not uncommon in the lighter forest.

*Notharcus pectoralis: Black-breasted Puffbird.

Not uncommon in the forest. Breeding: April 1935, in termite nest
(Skutch).

Notharcus tectus subtectus: Pied Puffbird.
Not uncommon in the forest near the laboratory.

*Malacoptila panamensis panamensis: White-whiskered Puffbird
(Panama Malacoptila).

Fairly common in the lighter forest, perching low. Breeding: In
the Canal Zone stub-tailed fledgling seen August 19, 1933 (Arbib).

Monasa morphoeus fidelis: White-fronted Nunbird (Goldman’s).
At least two seen April 6, 1950, on Wheeler trail (Wetmore).

Nonnula frontalis frontalis: Gray-cheeked Nunlet (Panama).
Seen in the forest September 7, 1935, off Drayton trail (D. W.

Lamm) ; seen in the forest, June 23, 1950, on Wheeler trail between
Barbour-Lathrop and Snyder-Molino trails (Stott).
Order PICIFORMES: Woodpeckers, Barbets, Toucans, and Allies
Family CAPITONIDAE: Barbets

Capito maculicoronatus maculicoronatus: Spotted-crowned Barbet.
Reported seen once in the forest (Sturgis).

NO. 5 BIRDS OF BARRO COLORADO ISLAND—-EISENMANN 31

Family RAMPHASTIDAE: Toucans

*Pteroglossus torquatus torquatus: Collared Aragari.

Common in the forest; seen almost daily around the clearing.
Notes: A high sneezy ksiyik, ksiyik (Skutch writes it pitit, pitit) ;
occasionally a soft nasal groan ang-uh; a staccato penk (Skutch).
Breeding: In natural hollow of a high horizontal branch, which was
first used as a sleeping hole by a group of six; incubation began late
March 1935; eggs hatched about April 11; 3 young emerged May
24, 26, 27 (Skutch).

*Ramphastos sulfuratus brevicarinatus: Keel-billed Toucan (Short-
keeled).

Very common in the forest ; seen daily around the clearing. Notes:
A dry, grunting or grinding werrk, werrk, werrk. The song is
described by Van Tyne as beginning with a hoarse low crr, crr, crr,
changing to cra, cra, cra, and finally settling to a shrill repeated cree,
cree, cree. Breeding: In natural cavities of large trees, eggs laid in
March and April (Van Tyne).

*Ramphastos swainsoni: Chestnut-mandibled Toucan (Swainson’s).

Common in the forest; seen almost daily around the clearing.
Notes: A loud, somewhat gull-like, yelping kee-you, tedick-tedick-
tedick ; sometimes kee-you, tedick.

Family PICIDAE: Woodpeckers

Celeus loricatus mentalis: Cinnamon Woodpecker (Fraser’s).

Occasionally seen in the forest (Chapman, Skutch, Stott).

*Centurus rubricapillus rubricapillus: Red-crowned Woodpecker
(Wagler’s).
Occasional in the clearing. A very common bird in more open
country. Notes: churr, churr; also wicka, wicka.

*Centurus pucherani pucherani: Black-cheeked Woodpecker (Pucher-
an’s).
Very common around the clearing. Notes: cherrr; also a higher
chirrirree, chirrirree. Breeding: December-June (Chapman, Eisen-
mann).

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, E17

*Dryocopus lineatus nuperus: Lineated Woodpecker (Panama Pile-
ated).
Common in the forest; often seen at the edge of the clearing.
Notes: A loud flickerlike wicka-wicka-wicka.

*Phloeoceastes melanoleucos malherbu: Crimson-crested Woodpecker
(Malherbe’s).
Fairly common in the forest ; often seen at the edge of the clearing.
Notes: A loud metallic cheerrr. Breeding: December-February ; young
left nest about February 20, 1925 (Chapman).

Order PASSERIFORMES: Perching Birds
Family DENDROCOLAPTIDAE: Woodcreepers 17

Dendrocolaptes certhia nigriorstris: Barred Woodcreeper.

Uncommon; seen December 24, 1934, in forest on Wheeler trail
(Davis) ; November 21, 1939 (Skutch) ; March 11, 1951 (two birds)
between Miller trail 2-4 (Collias) ; July 1, 1951, at northwest edge of
clearing (Eisenmann).

*Xiphorhynchus guttatus nanus: Buff-throated Woodcreeper (Law-
rence’s).

Common in the lighter woodland and forest edge. Notes: A series
of loud whistles, rising slightly in pitch and then dropping and slow-
ing down (Sturgis) ; a slurred cheer, a choe (Skutch). Breeding:
April 14, 1936, in partly covered box (Chapman) ; May 1935 in hole
in papaya tree (Skutch).

*Xiphorhynchus lachrymosus lachrymosus: Black-striped Wood-
creeper.
Not uncommon forest species; occasionally seen at the edge of the
clearing.

Campylorhamphus trochilirostris brevipenmis: Red-billed Sicklebill
(Venezuelan).
Seen August 17, 1935, on Snyder-Molino trail (D. W. Lamm).

17 This name, used by Chapman and Sutton, is adopted in place of the mis-
leading “woodhewer” of Sturgis and other authors.

Chapman (1938) lists Dendroplex picus “panamensis” (=extimus?) as
“Not common.” As this species has not been reported by any other observer,
and in Panama seems to be restricted to a narrow coastal strip along the Pacific,
some error is probable.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISEN MANN 33

*Glyphorynchus spirurus sublestus: Wedgebill.

Not uncommon in the forest. Notes: chif, chif. Breeding: July
12, 1949, nest in natural cavity, two fully-fledged young (Gross,
Loetscher ).

*Dendrocincla fuliginosa ridgwayi: Brown Woodcreeper (Dendro-
cincla).
Common in the forest. Notes: A loud squeaky sweeach or scheeah.

Family FURNARIIDAE: Ovenbirds and Allies

*Automolus ochrolaemus pallidigularis: Buff-throated Automolus
( Pale-throated).

A forest species of shrubbery and lower growth. Breeding: March
23, 1926, nest with two young in a clay bank (Van Tyne).

*Xenops minutus ridgwayi: Plain Xenops (Mexican).
Fairly common in the forest, generally feeds like titmouse among

outer branches of smaller trees and higher shrubs. Notes: tseet-tseet ;
a high cheep-cheep-cheep-cheep-cheep.

*Sclerurus guatemalensis guatemalensis: Scaly-throated Leafscraper
(Guatemalan Sclerurus).

Uncommon terrestrial forest species. Breeding: March 30, 1935,
with one nestling in burrow on stream bank near laboratory

(Skutch) .*8

Family FORMICARIIDAE: Antbirds

*Cymbilaimus lineatus fasciatus: Fasciated Antshrike.

A pair or two can generally be found at the north edge of the
clearing in the shrubbery or low trees. A bird of thickets in humid
areas. Notes: An easily imitated series of whistles, all on one note,
uttered at the rate of about two a second, and continuously repeated
for 8 to 10 seconds, cii, cii, cii, cii, cit or cwee, cwee, cwee, etc. The
alarm note is a nasal wasiurk.

18 The tawny-throated leafscraper (Mexican sclerurus)—Sclerurus me.i-
canus andinus—is included in Chapman’s 1929 (but not in his 1938) list. In
his later list, S. guatemalensis (omitted from the first list) is the only member
of the genus mentioned; hence the original listing was probably an error.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. BL

Thamnophilus doliatus nigricristatus: White-barred Antshrike (Black-
crested).

Occasional in the thickets around the clearing. Common in more
open country. Collected at Frijoles (Van Tyne). Notes: A rapid
whistled hooting, ending in a higher, more emphatic note hu-hu-hu-
hu-hu-hu-hu-hu-hu-hu-weng, rather similar to the analogous call of
the next species; a very harsh guttural garrr or ahrrr; a catlike whine.
Breeding: In the Canal Zone, fresh eggs July 17, August 12 (Har-
rower, Arbib).

*Thamnophilus punctatus atrinucha: Slaty Antshrike.

Most common antbird; a forest species. Notes: A rapid whistled
hooting, sometimes at a steady pace, but often accelerating into a roll,
ending in a nasal note, hu-hu-hu-hu-hu-hu-hu-hu-hu-hu-wenk; a
nasal wenk or wank, given alone; a note starting as a nasal and end-
ing in a growl, ang-grrrrr, or wang-grrrrr; a nasal, whining fiaah; a
harsh gr-gr-gr-gr-gr-gr. Breeding: Occupied nests December 22,
1930, January 7, 1937, February 14, 1935, March 17, 1935 (Skutch) ;
March 26, 1950 (Cottrell) ; May 7, 1935 (Skutch) ; July 4, 1949, July
6, 1950 (Eisenmann); July 5, 1947, July 30, 1947 (Mrs. Hobson) ;
July 13, 1949 (Gross) ; July 20, 1948 (Longenecker). All the July
nests contained eggs.

*Dysithamnus puncticeps puncticeps: Spotted-crowned Antvireo.
Not uncommon. A forest bird of low trees and shrubbery. Breed-

ing: Nests with two eggs, July 9, July 11, 1925, July 1949 (Gross) ;

food carried to nest near Shannon trail 1, April 22, 1951 (Collias).

Myrmotherula brachyura ignota: Pygmy Antwren.
Seen July 28, 1934 (Arbib).

Myrmotherula surinamensis pacifica: Streaked Antwren (Surinam).
Not common. A bird of the lower forest trees. Breeding: In the
Canal Zone, nest with eggs found July 13 (Stone, 1918).

*Myrmotherula fulviventris fulviventris: Fulvous-bellied Antwren
(Lawrence’s).
Very common in the more open part of the forest, frequenting
low trees and undergrowth. Notes: A squeaky queek, queek; a some-
what sweeter tseep, tseep; a rapid, continued series of chirps, syip-

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 35

syip-syip-syip; a series of high whistles going upscale, pii-peh-pey-
pih-piy-pee-pyee. Breeding: Nests with eggs, January 14, 1931, March
29, 1935 (Skutch) ; October 20, 1944 (Allen).

*M yrmotherula axillaris albigula: White-flanked Antwren (Black).

Very common in the forest, frequenting the lower branches of
smaller trees and the higher shrubbery, often in company with other
birds. Notes: A high piping on a descending scale, pee-a, peh-a, pi-a;
sometimes only peé-a, pii-a; also pee-pep-pip; also pip-pip or tit-tit ;
also psst, psst; also a dry trrrr or trtrtrir; a sharp whott; also a sweet
cheeup, cheeup. Breeding: April 3, 1935, full clutch; April 16, 1935,
very young nestling ; May 7, 1935, eggs (Skutch).

*Microrhopias quixensis virgata: Dotted-winged Antwren (Panama).

Common in the more open parts of the forest, frequenting the lower
branches of smaller trees and the higher shrubbery. Notes: wit-wit or
tit-tit; also a higher cheet, cheet; a very dry thrrrr; a very high chick-
like cheep, cheep, cheep, varied by a keeya resembling a note of
Myiozetetes. Breeding: Nest building, February 22, 1935 (Skutch).

*Cercomacra tyrannina rufiventris: Dusky Antbird (Tyrannine).

Common in thickets and undergrowth of forest openings and edge.
Notes: A song of five to eight whistled notes, going upscale but not
smoothly, with pauses after the first two notes, the remainder given
very rapidly and rising in pitch, pii, pii, pi-pi-pi, or pii, peh, pipipt-
peepee. A call, teé-o0, teé-oo. Breeding: Nest, April 29, 1935
(Skutch).

Myrmeciza longipes panamensis: White-bellied Antbird.

Not uncommon, in second-growth woodland, usually on the ground.
Notes: A “loud rail-like whinny” (Chapman).

*Myrmeciza exsul exsul: Chestnut-backed Antbird (Sclater’s).

Common in the forest and forest edge; frequents thickets, keep-
ing low, but is not terrestrial. Notes: A loud, emphatic, easily imi-
tated whistle, deh, deé-a, or deh, deh, deé-a, which Chapman para-
phrases as “Come here” and “Come right here.” Also a clucking
quick-ick, and a nasal, catlike nyaah. Breeding: Nests with eggs, July
13, 1927, July 25 and August 5, 1925 (Gross).

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

*Formicarius analis panamensis: Black-faced Antthrush (Panama).

Common terrestrial forest species; much more often seen than
heard. Notes: A phrase of three (occasionally four) deliberate plain-
tive whistles, the first longer and about a half tone higher than the suc-
ceeding notes, which are on the same pitch. The bird will decoy to an
imitation of this call. The alarm note is “a sharp, clear, some-
what explosive fleet, sometimes repeated in a continuous sequence”
(Skutch). Breeding: Nest with one egg in tree stump, May 19, 1929
(Cleaves).

*Gymnopithys leucaspis bicolor: Bicolored Antbird.

Fairly common in the forest, on or near the ground; often with
army ants. Notes: A low, rather whining chirrrrr. Breeding: Nest
with eggs, April 15, 1927 (Van Tyne, 1944).

*Hylophylax naevioides naevioides: Spotted Antbird.

Common in forest undergrowth, generally off the ground, usually
with army ants. Notes: Most characteristic is a rather soft, deliberate,
oft-repeated teé-de, teé-de, teé-de, teé-de, teé-de, gradually slowing
in time and slightly descending in pitch. Also a sharp, whistled supeé,
pee, pee, pipipipi, the last group of notes very fast; also a low chirrr
or tsirrr. Skutch mentions as additional alarm notes a high sharp
psip and a short soft peep. Breeding: Nests with eggs, April 18, May
7, May 16, 1935 (Skutch), July 16, 1925, July 1949 (Gross) ; June
24, 1951, hatched July 2 (Milne, Laughlin, Eisenmann).

*Phaenostictus mcleannan mcleannant: Ocellated Antthrush (Mc-
Leannan’s).

Not uncommon in the forest; a terrestrial species most often seen
with army ants. Notes: A soft, very fast series of 5 to 8 whistles,
slightly upscale, puhpiipehpeypihpee; a sharp wheerrr or cheerrr.
Breeding: July 10, 1950, adults feeding stub-tailed fledgling (Eisen-
mann).

Grallaria perspicillata perspicillata: Streak-chested Antpitta (Law-
rence’s).

Fairly common terrestrial forest species; more heard than seen;
frequently heard from the laboratory. Notes: A series of clear,
melancholy whistles, at first rising a little in pitch, then falling off
in three descending couplets, deh, dee, dee, dee, dee, dee, deé-eh, déh-
oh, doh-a. The notes preceding the couplets vary in number, but the

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN SV

couplets are almost always three. There is usually a few minutes’
pause before repetition of the song; by imitation of it the bird can
be decoyed into view.

Family COTINGIDAE: Cotingas

Cotinga nattererti: Blue Cotinga (Natterer’s).

A tree-top forest species ; probably not uncommon, but infrequently
seen; occasionally noted in the tall trees at the edge of the clearing.
Breeding: February-March, 1927, 1928 (Chapman).

* Attila spadiceus sclateri: Polymorphic Attila (Sclater’s).

An arboreal bird of the lighter woodland, often perching fairly
low. Notes: A “measured, emphatic beat-it, beat-it, beat-it, no-6-w,”
also a sharp twitter in flight (Chapman).

*Laniocera rufescens rufescens: Speckled Mourner.

“Rare” (Chapman). Adult female collected March 3, 1926 (Van
Tyne). A little-known forest species. Resembles the next species
in size (length, 7.8 inches) and color, being rufous above and tawny
below ; but differs in having the wing coverts dusky, speckled with a
rufous; a tuft of yellow feathers on the sides of the breast (usually
concealed), and very indistinct barring on the chest.

*Rhytipterna holerythra holerythra: Rufous Mourner (Rufous Lip-
augus).
“Not common” (Chapman). Collected April 25, 1926 (Van Tyne).
An arboreal forest species.

*Lipaugus unirufus castaneotinctus: Rufous Piha (Panama Lathria).

Rather common arboreal forest species. Notes: A very loud, em-
phatic, whistled chu-weé-oo, repeated from time to time. Chapman
describes it as a “staccato, explosive see-you I-see-you, a long sweep-
ing silvery whistle.”

Pachyramphus cinnamomeus cinnamomeus: Cinnamon Becard.
An infrequently noted arboreal forest species.

Pachyramphus polychopterus cinereiventris: Black-backed Becard
(Gray-bellied).
“Infrequent” (Chapman). An arboreal bird of open woodland
clearings. Breeding: In the Canal Zone, nest with eggs found June
18 (Stone, 1918).

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Tityra semifasciata costaricensis: Masked Tityra (Costa Rican).
Common arboreal species, especially at the edge of forest clearings.
Notes: A dry quert, quert. Breeding: May 28, 1935, nest with eggs
in natural tree cavity (Skutch); June 29, 1948, feeding young in
woodpecker hole (Eisenmann) ; June 28, 1951, feeding young (Ellis).

*Erator inquisitor fraserii: Black-crowned Erator or Tityra
(Fraser’s).
Not uncommon arboreal forest species; occasional at the edge of
the clearing. Notes: Like those of the preceding species, but “drier
and less ‘grunty’” (Skutch).

*Querula purpurata: Purple-throated Fruit-crow.

Very common, noisy, arboreal, forest species, wandering about
in small bands; frequently heard from the laboratory. Notes: A
variety of loud, often liquid or tremulous cawing calls, kweé-a-wuh,
kweé-a-wuh; kwoo, kwoo; kwék-kwék-oo; cherkaw, cherkaw; kwek-
kwek. Breeding: Pair noted building nest at northwestern edge of
clearing June 28, 1951; incubation apparently begun July 14 (Ellis,
Eisenmann).

Family PIPRIDAE: Manakins

*Pipra mentalis minor: Red-capped or Yellow-thighed Manakin.

Common in the lower levels of the lighter forest. Notes: sick,
seceeeceeee, the second note very high, thin, and rising in pitch; also
a tsick or seck, or psit, which Skutch writes is sometimes repeated.
The male likewise produces a snap and whirring, doubtless with its
wings. Breeding: Nests with eggs, 3 in March (earliest March Io),
2in April, 1 May 7 (Skutch) ; July 8, 1947; young left July 29 (Mrs.
Hobson) ; July 7, 1950 (Mrs. F. Ryan, Eisenmann).

*Manacus vitellinus vitellinus: Golden-collared Manakin (Gould’s).

Common in the lighter second-growth woodland, particularly near
the lake shore, keeping low. Notes: Vocal notes are a whistled peé-
you, and a softer chee-pod or chee-wodoo. Other sounds, probably
made by the wings, are a loud snap, reminiscent of stones struck
together ; a reedy whir; a ripping sound like the tearing of cloth; and
a less loud snip (Chapman 1935; Arbib). Breeding: Nests with eggs
February 27-August 1 (Skutch, Van Tyne) ; July 22, 1925, July 25,
1925 (Gross). In the Canal Zone, off the island, nests with eggs have
been found in March, April, May, June, and August (Stone, 1918,
Harrower, Arbib).

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 39

Chiroxiphia lanceolata: Lance-tailed Manakin (Sharp-tailed).

Female seen March 11, 1946, at Fred Miller trail 15 (Wetmore).
A bird of scrub and second-growth thickets, common on the Pacific
slope of Panama. Notes: The basic note is a mellow, but rather
hollow, doh, with a bell-like quality, which is repeated alone, or
rapidly reiterated do-do-do-do-do, or accelerated into a musical roll
drrrrrrrr, or varied into déwee, déwee-oh, and deé-o. Another more
elaborate phrase goes whit, coo-cod, do-do-do-do-do-whit, and still
another, with a somewhat oriole quality, do-do-dee-do-deh. There is
also a very different nasal whang. Breeding: In the Canal Zone and
vicinity, nests with eggs have been found on August 11 and September
13 (Hallinan).

Schiffornis turdinus panamensis: Thrushlike Manakin.

“Rare” (Chapman). A forest bird of the undergrowth. Rather
large (length, 6.5 inches) for a manakin. Above plain brown; below
cinnamon on throat and chest, pale olive on breast and flanks, fading
to yellowish on abdomen. Immature birds are more uniformly cinna-
mon below, paler on throat, and grayish on abdomen.

Family TYRANNIDAE: Tyrant Flycatchers

*Tyrannus tyrannus: Eastern Kingbird.

Common migrant; sometimes noted in numbers; March 22, 1927,
30 seen (Chapman) ; collected April 21, 27, 1926, seen April 12,
April 17, 1927 (Van Tyne). May 6, 1935, at least 100 (Skutch) ;
September 14-16, 1938, 6 to 8 together (Eisenmann). Recorded in
Panama March-May and August-October.

*Tyrannus melancholicus chloronotus: Tropical Kingbird (Lichten-
stein’s).
Common in the clearing and along the lake shore. Notes: A rapid
twittering piririri, sometimes piriri, also feet-feet-feet. Breeding:
Nests February-June (Chapman, Skutch).

Tyrannus donunicensis dominicensis: Gray Kingbird.

Seen February 1940 (Davis). Migrant and winter visitant in Pan-
ama November-April.

*Legatus leucophaius leucophaius: Piratic Flycatcher (Striped).

Pairs occur around clearings and lake shore, and elsewhere in
breeding colonies of victim species. Notes: A monotonously repeated

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

wee-yee? piririree ; often the wee-yee? is given alone. Breeding: Seizes
newly built closed nests of other species that are harried until
they desert. Appropriating and apparently using nests in oropéndola
colony February-April (Chapman, Sturgis) ; appropriating Myioze-
tetes nests in late March (Chapman).

*Myiodynastes maculatus difficilis: Streaked Flycatcher (Noble).
One or two pairs frequent the laboratory clearing and others occur
along the lake shore. Notes: A loud heavy chup or check ; also eéchup ;
also chupeé or chupeét; also, by the male, chup, weéarooweép, the
latter phrase given after a slight pause and of such different, rather
musical though squeaky quality that it seems as if made by another
bird. Breeding: Builds nests in tree holes, roof corners, boxes, and
other natural or artificial crannies and hollows. Late December 1925,
woodpecker hole (Chapman) ; March 17, 1950, incubating (Cottrell) ;
April 6, 1950, angle of roof, April 24, 1947, angle of roof (Wet-
more) ; April 1935, bird box, May 1935, woodpecker hole (Skutch) ;
June 29, 1948, feeding young in woodpecker hole, June 27, 1950,
incubating in angle of roof, young fed July 7 (Eisenmann) ; July
I, 1947, feeding young in angle of roof (Mrs. Hobson) ; July 7-10,
1949, eggs laid, young hatched July 25, in angle of roof (Gross).

*Megarhynchus pitangua mexicanus: Boat-billed Flycatcher.

A pair can generally be found in the laboratory clearing and others
are well distributed along the lake shore. Notes: A harsh, rattling
keerrrrrik-keé several times repeated; also krrrrreek. Breeding:
June 27, 1925, nest with large young (Van Tyne); July 15, 1947,
feeding young outside nest (Mrs. Hobson) ; June 28—July 3, 1951,
incubating (Eisenmann). In the Canal Zone young have been found
in the nest June 2 (Stone, 1918).

*Myiozetetes cayanensis harterti: Rusty-margined Flycatcher (Cay-
enne).

A pair or two can often be found in the laboratory clearing ; others
are frequent along the lake shore and are also occasionally seen in
the treetops in the forest. Notes: A rather high fee-ee; also feéee-you.
Breeding: January-May (Chapman, Skutch). In Panama City found
incubating in early July (Eisenmann).

*Myiozetetes similis columbianus: Vermilion-crowned or Social Fly-
catcher (Colombian).

A couple of pairs can usually be found in the clearing and birds

are common along the lake. Notes: The common call is a loud, harsh

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 4I

keé-you or kreé-you, also a longer keé-you, kee-kee-kee-kee-kee.
Breeding: Vebruary-June (Chapman, Skutch, Van Tyne), with egg-
laying chiefly March-May.

Myiozetetes granadensis granadensis: Gray-capped Flycatcher.

Occasional along the lake shore. Notes: A heavy chip; a loud,
harsh kree-you. Breeding: In the Canal Zone, nests with eggs have
been found in April and May (Hallinan).

*Pitangus lictor panamensis: Lesser Kiskadee (Lictor Flycatcher).

Not uncommon in the coves of Gigante Bay and around the mouth
of Fuertes Estero. Notes: Buzzy and squeaky, dzai; also dewey, and
dzee; also an excited dzai-dey-dzéy-dah repeated over and over ; also
dzey-ee-dey. Breeding: In the Canal Zone, female with egg July 9
(Stone, 1918).

*Myiarchus crinitus: Great-crested Flycatcher.

Migrant and winter visitant. Collected March 19, 1927, and May
3, 1926 (Van Tyne). Recorded in Panama from October 6-May 3.

*Myiarchus ferox panamensis: Short-crested Flycatcher (Panama).

Fairly common in the clearing and in the lighter woodland, usually
sitting rather low. Notes: A rather soft, whistled whee; also a whis-
tled whee-hee-hee-hee-hee-hee-hee-hee-hee.

*Myiarchus tuberculifer brunneiceps: Dusky-capped Flycatcher
(Black-crested).
Fairly common at the edge of the clearing and in the lighter wood-
land. Notes: A loud, throaty, whistled wheeeew, with a bit of r in
it; also a whistled wheeeep, suggesting the great crested flycatcher.

*Contopus virens: Eastern Wood Pewee.

Migrant. Collected April 23, 1926, April 29, 1927, and May 14,
1927 (Van Tyne). Seen and heard calling February 9, 10, 1940,
at edge of clearing (Mrs. Fry and party). In Panama this species
has been taken April 17-May 14 and September 8-November 8. Birds
of this genus were noted by the writer in September 1938. As both
C. virens and the western C. richardsoni migrate through the Canal
Zone, sight identifications of silent birds are unreliable. The difficulty
is further increased by the possible occurrence of the tropical pewee
(Contopus cinereus brachytarsus), which is a common Canal Zone
resident.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

Empidonax flaviventris: Yellow-bellied Flycatcher.
Seen October 17, 1944 (Allen). A winter visitant, recorded in
Panama September 8-April 14.

*Empidonax virescens: Acadian Flycatcher.
Collected February 19, 1927 (Chapman). A winter visitant, re-
corded in Panama October 16-April 7.

Empidonax traillu: Traill’s Flycatcher.

“Winter visitant ; no specimens” (Chapman). Both the eastern and
western forms of this species appear to winter in (and at least regu-
larly migrate through) Panama; records for traillii: August 21-May
13; for brewstert: September 17-May 9.*°

*T erenotriccus erythrurus fulvigularis: Ruddy-tailed Flycatcher (Ful-
vous-throated).
Not uncommon in the forest. Notes: A thin tseé-oo-tee. Breeding:
March 1935, two nests with eggs; May 18, 1935, nest with nestlings
(Skutch).

*Myiobius sulphureipygius aureatus: Sulphur-rumped Flycatcher
(Myiobius).
Not uncommon along woodland streams. Breeding: Nests hung

over stream at the rear of the clearing, March and April 1935
(Skutch).

*M yiobius atricaudus atricaudus: Black-tailed Flycatcher (Myiobius).

Fairly common along streams and the lake shore. Breeding: Nests
hung low over water; nests with eggs June 28, July 8, July 9, July
10, August 9, 1925 (Gross).

*Onychorhynchus mexicanus fraterculus: Northern Royal Flycatcher
(Colombian).
Uncommon woodland bird, favoring the vicinity of water. Breed-
ing: Nesting over stream February-March, 1935 (Skutch).

*Platyrinchus coronatus superciliaris: Golden-crowned Spadebill
(Lawrence’s).

Occasionally noted in the forest, sitting in low trees or shrubs.

19 The least flycatcher (E. minimus) has been taken at least once in the
Canal Zone.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 43

*Cnipodectes subbrunneus panamensis: Brownish Flycatcher.

Occasionally noted in the forest. Breeding: April 5, 1935, seen
building nest (Skutch).

Tolmomyias sulphurescens flavo-olivaceus: Sulphury Flycatcher (Yel-
low-olive).
Seen May 22, 1933 (Carleton). A bird of light woodland and
forest edge. Breeding: In the Canal Zone, nest with young found
May 28 (Stone, 1918).

Tolmomyias assimilis flavotectus: Yellow-margined Flycatcher.

Uncommon in the forest. Breeding: April 6, 1948, building nest
(Wetmore).

*Rhynchocyclus olivaceus bardus: Olivaceous Flatbill (Equinoctial
Flycatcher ).
Not uncommon in second-growth woodland and forest edge. Breed-
ing: April 13, 1937, young about four days old in nest (Gilliard).
Four sleeping nests found in February and March, 1935 (Skutch).

Todirostrum cinereum finitimum: Common Tody-flycatcher (North-
ern).

Occasional in the clearing; a bird of open areas with scattered
trees and thickets. Notes: A short srrrr; a sharp chip. Breeding:
April 1937 (Chapman) ; 2 eggs, May 2, 1935 (Skutch) ; nest with
2 eggs, hatched July 10 (Eisenmann).

Todirostrum sylvia schistaceiceps: Slate-headed Tody-flycatcher.
“Not common” (Chapman). A bird of low shrubbery in clearings
and open country.

*Oncostoma olivaceum: Southern Bentbill (Lawrence’s Bent-billed
Flycatcher).
Not uncommon in thickets in the lighter forest. Breeding: Nests,
April 1937 (Chapman) ; two eggs, May 2, 1935 (Skutch). In the
Canal Zone, nest with eggs, August 7 (Harrower).

Capsiempis flaveola semiflava: Yellow Flycatcher.
Pair seen in the clearing November 1935 (Skutch). A shrub bird
of open country.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

*Elaenia flavogaster pallididorsalis: Yellow-bellied Elaenia (North-
ern).

Male collected April 21, 1927, on Slothia Island in the laboratory
cove (Van Tyne). A very common bird of open country and clear-
ings. Notes: A loud, harsh krreéup, weekrreéup; sometimes only
krreéup or krreep; also wrrree. Breeding: In the Canal Zone, occu-
pied nests found March 31 (Hallinan) ; July 21 (Harrower) ; but
mostly in April-June (Harrower ; Stone, 1918; Eisenmann).

Elaenia chiriquensis chiriquensis: Lesser Elaenia (Lawrence’s).

Occasional in the clearing ; an open-country bird. Notes: A wheece
or feeee, sometimes slightly trilled freeee; also feé-o0; also a buzzy
peb-zii. Breeding: In the Canal Zone and nearby Panama, June 18,
juvenals being fed by adults; June 24, nest with young a few days
old (Eisenmann) ; July 17, nest with incubation apparently going on
(Arbib) ; July 30, nest with young (Imhof).

Sublegatus arenarum atrirostris: Scrub Flycatcher (Smooth).
“Not uncommon” (Chapman). A bird of scrubby country; com-

mon on the Pacific slope of Panama. Breeding: In the Canal Zone
in April (Imhof).

Camptostoma obsoletum flaviventre: Southern Beardless Flycatcher
(Yellow-bellied Camptostoma).
Occasionally noted in the clearing; a bird of open-country thickets
and second growth. Breeding: March 7, 1949, feeding full-grown
young near laboratory (Wetmore).

Tyranniscus vilissimus parvus: Paltry Tyrannulet (Lesser Paltry
Flycatcher).

Common around the edge of the clearing and observed at various
places on the lake shore and in the forest. Notes: pee-ee-yip or chee-
yip, frequently uttered. Not listed by Chapman and not collected,
but noted in recent years by all experienced observers.

Tyrannulus elatus panamensis: Yellow-crowned Tyrannulet.

Common in the small trees of the clearing, and noted at various
places on the lake shore and at Fuertes House. Notes: A clear, fre-
quently given whistle, deé-ee, deéa. Breeding: July 3, 1950, adults
feeding a stub-tailed juvenal (Eisenmann). Not listed by Chapman
and not collected, but noted in recent years by most visitors.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 45

Microtriccus brunneicapillus brunneicapillus: Brown-capped Tyran-
nulet.

Fairly common in the clearing. Notes: A high, piping whistle
usually of five notes, descending in pitch, pee-pih-pey-peh-pii, and
uttered so fast that the whole phrase lasts only about a second; also
a single high, clear peep. Not listed by Chapman and not collected.
I have regularly noted it around the laboratory on my many visits
in June, July, and August.

*Pipromorpha oleaginea parca: Ochre-bellied Flycatcher (Bang’s
Pipromorpha).

Rather common woodland species, particularly in low trees of hill-
sides and ravines. Breeding: March 2, 1926, 2 nests with eggs hang-
ing against large tree on Wheeler trail, eggs of one nest hatched
March 25, 1926; July 12, 1925, 2 nests hanging against cut-bank of
Lutz Stream, eggs of one nest hatched July 15, 1925 (Van Tyne) ;
November 23, 1939, nest (Skutch). In the Canal Zone, nest with
3 eggs, May 14 (Stone, 1918).

Family HIRUNDINIDAE: Swallows

*Iridoprocne albilinea: Mangrove Swallow.

Common, flying low over the lake. Breeding: Probably nests in
holes in partly submerged tree trunks of the drowned forest.

Riparia riparia riparia: Bank Swallow.

Migrant; four seen April 17, 1942, with migrating barn swallows
(Imhof). Reported in the Canal Zone February 22-April 17, August
26-November 30.

Stelgidopieryx ruficollis uropygialis: Rough-winged Swallow (Pan-
ama).
Fairly common, flying over the lake, sometimes over the clearing.
The northern race serripennis has been taken in winter in the Canal
Zone.

*Hirundo rustica erythrogaster: Barn Swallow.

Common migrant; March 23 (Chapman)—May 17 (Van Tyne) ;
occasional in winter, December 30, 1934 (Davis). Migration in the
Canal Zone is chiefly from March to May and August to October.

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

*Progne chalybea chalybea: Gray-breasted Martin.

Very common, flying over the lake and the clearing and often
perching on trees along the shore. Breeding: April 6, 1950 (Wet-
more) ; May 20, 1935 (Skutch) ; July 2, 1951, nestling at nest near
Slothia Island (Laughlin) ; all three nests in tin-can channel markers.

Phaeoprogne tapera: Brown-chested Martin.

July 4, 1949, three recognized flying over the laboratory were
later studied at close range, perched on a tree near the lake with the
preceding species (Eisenmann), and were shown to Dr. A. O. Gross.
At least 12 were noted perched on wires at Frijoles in July 3, 1951
(Eisenmann). Though never previously reported from Panama, this
South American species was regularly seen by me about the city of
Panama and in the Canal Zone during late June and July, 1949 and
1951, often perched on wires in numbers up to about 100. The race
of southern South America (fusca) is known to be migratory. This
species, though very much larger (length, 6.5 inches), closely resem-
bles the bank swallow in color and pattern. The birds seen had notice-
able speckling below the brown breastband, thus suggesting the south-
ern fusca.

Family CORVIDAE: Jays, Crows, and Allies

*Cyanocorax affinis zeledoni: Black-chested Jay (Talamanca).

Small groups are occasionally noted in the forest. Notes: A loud
metallic chung, chung; also a harsh wahk; also a rapid, dry, and loud
chikidik ; also a loud kiyiytying.

Family TROGLODYTIDAE: Wrens

*Thryothorus castaneus castaneus: Bay Wren.

“Rare” (Chapman). Collected March 22, 1926 (Van Tyne). A
bird of thickets in most woodland and along stream edges. Breeding:
In the Canal Zone, nest with egg found July 28 (Stone, 1918).

Thryothorus leucotis galbraithu: Buff-breasted Wren (Galbraith’s).

Occasional in thickets around the clearing. A common species
(collected at Frijoles) of moist thickets, in fairly open areas and the
edge of clearings.”

20 The plain wren (7. modestus elutus), common in larger clearings, may
be expected to occur. It lacks the sharp black wing barring and is less buffy on
the breast.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 47

Thryothorus fasciato-ventris albigularis: Black-bellied Wren.
Occasional in the thickets at the edge of the clearing.

*Troglodytes musculus inquietus: Southern House Wren (Panama).

A pair generally can be found around the laboratory and sometimes
at one of the houses along the shore of the island. Notes: Most of
them sufficiently resemble those of the northern 7. aedon to be readily
recognized. Breeding: Egg dates, early January through August
(Chapman, Van Tyne, Skutch, Eisenmann).

*Henicorhina leucosticta pittiert: White-breasted Wood Wren (Sclat-
er’s).
Uncommon ; collected by Van Tyne (who identifies it as this race)
and also by Chapman. A bird of forest undergrowth and fallen treetop
tangles.

*Cyphorhinus phaeocephalus lawrencu: Song Wren (Lawrence’s
Musician).

Fairly common in the forest in thickets, occasional at the edge of
the clearing. Notes: Call, a guttural ciutta, cuitta, cuitta. Song, a variety
of whistles usually intermixed with cutta notes, per-cutta wheet, per-
cutta whoot, per-cutta wheet, per-cutta whoot; per-cutta-cuttle wheet
whoot; per-cutta whoot whoot (Harrower). Two sleeping nests with
four or five occupants February-March 1935 (Skutch).

*Microcerculus philomela philomela: Nightingale Wren.

A little-known forest species. Collected March 23, 1926 (Van
Tyne). Seen December 30, 1930 (Chapman, Skutch). May 14-23,
1933, a total of six found singing in various parts of the island
(Carleton).

Family MIMIDAE: Mockingbirds and Thrashers

Dumetella carolinensis: Catbird.
“Rare winter visitant” (Chapman). Recorded from Panama
October 24-April 23.

Family TURDIDAE: Thrushes

*Turdus grayt casius: Clay-colored Robin (Bonaparte’s).
A pair or two can generally be found in the clearing. Notes: Its
song and most of its calls are much like those of the American robin

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

(T. migratorius), but it has a distinctive, whining call, often inter-
minably repeated, sounding like chee-weé-va or kee-eé-wuh, sometimes
wayewoo. Breeding: Egg dates in the Canal Zone and adjacent Pan-
ama range from February-August (Hallinan, Harrower, Eisenmann ;
Stone, 1918).

Hylocichla mustelina: Wood Thrush.
Rare winter visitant ; seen December 29, 1926 (Chapman), March
21, 1935 (Skutch).

*Hylocichla ustulata swainsonti: Swainson’s Thrush (Olive-backed).

Migrant. Collected April 17, 1927 (Van Tyne), seen April 4, 5, 6,
1935 (Skutch), April 17, 1942 (Imhof), October 14, 1944 (Allen).
Fairly common migrant through Panama October 7-November 9 and
March 17-May I.

Family SYLVIIDAE: Warblers, Gnatcatchers, and their Allies

*Polioptila plumbea bilineata: Tropical Gnatcatcher (Lawrence’s).

Not common; a bird of light second-growth and scrubby areas.
Notes: A thin nasal tzeet, tzeet; a nasal twang.

*Ramphocaenus melanurus rufiventris: Long-billed Gnatwren (Ant-
wren).
Fairly common in thickets in the lighter woods. Breeding: In Pan-
ama City, nest with two newly hatched young, July 15, 1950 (Eisen-
mann).

Family CYCLARHIDAE: Pepper-shrikes

Cyclarhis gujanensis: Rufous-browed Pepper-shrike.

Seen singing March 9-10, 1950, near the laboratory (Wetmore).
Previously unreported from the Canal Zone, and known in Panama
only from open woodland and scrub on the Pacific slope. Length, 5.6
inches. Above olive-green, with brownish cap, rufous superciliaries,
gray sides of head ; below largely yellow, fading to white on abdomen.

Family VIREOLANIIDAE: Shrike-vireos

*Smaragdolanius pulchellus viridiceps: Green Shrike-vireo.

A forest bird of the treetops, very hard to see, but probably not
uncommon. Notes: Whistles a monotonous “ ‘one-two three’, all on
the same note, with a tireless persistence” (Chapman). Length, 5.3

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 49

inches. Entirely brilliant green, with a yellow throat and blue hind-
neck.

Family VIREONIDAE: Vireos

Vireo flavifrons: Yellow-throated Vireo.

Seen March 12, 1935 (Skutch). Uncommon winter visitant in
Panama, November 3-March 12.

Vireo olivaceus: Red-eyed Vireo.
“Migrant” (Chapman). A common migrant and occasional winter

visitant in Panama, August 14-May 1, but chiefly seen in September
and October, March and April.

Vireo flavoviridis flavoviridis: Yellow-green Vireo.

Very common in the treetops in the forest and at the edge of the
clearing. Panama-breeding birds are sometimes separated under the
name insulanus. Notes: Calls are like those of V. olivaceus; song is
also similar, but the phrases are shorter, drier, and more monotonous,
and the utterance is less continuous. Breeding: In the Canal Zone,
March 26 (Imhof).

*Hylophilus decurtatus pusillus: Gray-headed Greenlet (Pachysylvia).

Not uncommon in the lighter woods, generally moving in small
groups among the lower trees; sometimes seen at the edge of the
clearing. Notes: A rapid, emphatic, whistled cheetsacheét, repeated
at intervals of about 5 seconds; sometimes the note sounds more like
itsachéyit. Breeding: In the Canal Zone, young collected June 25
(Stone, 1918).

Family COEREBIDAE: Honeycreepers

*Coereba flaveola columbiana: Bananaquit.

Common in the clearing among the smaller trees and flowering
shrubs. Notes: A buzzy, high, unmusical zizi-zizi-zizt-zizi-ziz, some-
times accelerating to zizizizi. Breeding: Nests, January 20, 1935,
February-April 1935 (Skutch) ; August 30, 1947 (Eisenmann) ; No-
vember 20, 1939 (building), December 29, 1930 (Skutch). Old nests
are often used for sleeping, and sleeping nests are also built.

*Dacnis cayana ultramarina: Blue Dacnis (Ultramarine).
Common in and around the clearing and trees of the lake shore.

Notes: tseet. Breeding: Two juvenals being fed July 28, 1947 (C. A.
Moore).

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Dacnis venusta venusta: Scarlet-thighed Dacnis.

Seen February 8, 1951, 2 males and 2 females at Fuertes House
(Collias) ; June 26, 27, 1951, male and female at south edge of labora-
tory clearing (Eisenmann). A species rarely recorded in the Canal
Zone. Length, 4.75 inches. Male: Turquoise blue above, including
sides of head; black forehead, wings, tail, and underparts; thighs
scarlet. Female: Olive above; bluish-green sides of head and neck;
buffy underparts ; thighs tinged with red.

*Cyanerpes cyaneus carneipes: Blue Honeycreeper.

Usually very common in and around the clearing—at least in June
and July; 20 individuals (mostly adult males) on June 27, 1951, in
one tree; not noted in August 1945, September 1938 (Eisenmann),
or in October 1943 (Allen). Notes: st, st or tsst, tsst; also ggee.
Breeding: Feeding juvenal, April 6, 1937 (Gilliard).

Cyanerpes lucidus isthmicus: Shining Honeycreeper.

July 1, 2, 3, 6, 11, 1950, I to 3 individuals of both sexes seen at
close range in and around the clearing (Eisenmann) ; March 17, 1951,
pair (Scholes). Length, 4 inches. Male: General color deep blue, be-
coming azure on the crown; black throat, wings, and tail; legs bright
yellow. Female: Greenish above, becoming grayish-blue on the head ;
underparts whitish, the chest streaked with blue, the sides with green-
ish; legs greenish.

*Chlorophanes spiza arguta: Green Honeycreeper.

Common in and around the clearing, particularly in the tall trees.
Notes: pst, pst; also a short nasal grunt uhr, given by the male. Breed-
ing: July 7, 1950, female gathering nesting material (Eisenmann).

Family PARULIDAE: Wood Warblers

Mniotilia varia: Black-and-white Warbler.

Fairly common winter visitant; late date, March 18, 1933 (Carle-
ton). Recorded in Panama August 24-March 26.

*Protonotaria citrea: Prothonotary Warbler.

Fairly common winter visitant, especially along the lake shore;
November 24, 1939 (Skutch)—March 14, 1933 (Carleton). Recorded
in Panama August 1-March 14.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 5I

Helmitheros vermivorus: Worm-eating Warbler.

Seen February 26, 1935 (Skutch). A winter visitant recorded in
Panama December 15—March 16.

Vermivora chrysoptera: Golden-winged Warbler.

Winter visitant (Chapman); September 19, 1951 (Bourliere).
Recorded in Panama October 24—April 16.

Vermivora peregrina: Tennessee Warbler.

Fairly common migrant and winter visitant; October 23, 1944
(Allen)—March 24, 1950 (Cottrell). Recorded in Panama October
17—April I.

*Dendroica petechia aestiva: Yellow Warbler.

Common winter visitant. Collected April 17, 1927, May 4, 1926
(Van Tyne). Seen September 12, 1938 (Eisenmann)—April 23,
1935. Recorded in Panama August 17—May 4. Barro Colorado sight
records may not all be this subspecies, but they relate to the northern
birds treated until recently as the species D. aestiva.

*Dendroica pensylvanica: Chestnut-sided Warbler.

Common winter visitant, October 25, 1944 (Allen)—April 15, 1935
(Skutch). Recorded in Panama September 22—April 15.

Dendroica fusca: Blackburnian Warbler.

Seen March 20, 1950 (Cottrell). Recorded in Panama as migrant
September 27—-November 17, March 5-April 28.

*Dendroica castanea: Bay-breasted Warbler.

Fairly common migrant and winter visitant, October 22, 1944 (Al-
len)—April 5, 1927 (Van Tyne). Recorded in Panama October 22-
April 27.71

21The blackpoll warbler (Dendroica striata) is the subject of two sight
reports, which are remarkable as this species migrates through the West Indies
and has apparently never been recorded in Central America. Dr. A. A. Allen
writes that November 6, 1944, he observed one “close enough so that I had no
trouble seeing the light-colored legs and white under tail coverts.’ Mrs. G. G.
Fry writes that on February 8 and 9, 1940, she heard a song, which she recog-
nized as that of this species, and that one of the members of her party, Mrs.
L. J. Francke, saw the bird and said it was a male in spring plumage.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Seiurus aurocapillus: Ovenbird.

Winter visitant. Reported singing February 19, 20, 1940 (Mrs.
Fry). Late date, March 18, 1933 eaticton): Recorded in Panama
November 26—April 15.

*Seiurus noveboracensis notabilis: Small-billed Waterthrush (North-
ern).

Common winter visitant along forest streams and lake shore. Six
specimens taken by Van Tyne March 11, 1926—May 4, 1926, April
17, 1927, are identified by him as this race. Other forms probably
occur. Seen October 25, 1944 (Allen)—May 4, 1926 (Van Tyne).
Recorded in Panama September 18—May 4.

*Seiurus motacilla: Louisiana Waterthrush.

Collected March 12, 1926 (Van Tyne). Recorded in Panama Au-
gust 24—March 18.

Oporornis formosus: Kentucky Warbler.

Uncommon winter visitant. Late date, March 28, 1935 (Skutch).
Recorded in Panama September 8—March 28.

Oporornis philadelphia: Mourning Warbler.

Uncommon winter visitant (Chapman, Skutch). Recorded in Pan-
ama September 24—April 14.??

Wilsonia citrina: Hooded Warbler.

Male seen September 24, 1951 (Bourliere). Rare migrant and
winter visitant in Panama.

Wilsoma canadensis: Canada Warbler.

Migrant, September 24, 1951 (Bourliere); October 20, 1944
(Allen) ; March 25, 1950 (Cottrell) ; April 16, 20, 1935 (Skutch).
Recorded in Panama September 21-October 20, March 25—May 19;
sometimes winters.

Setophaga ruticilla: American Redstart.

Winter visitant, January 18, 1931 (Skutch), February 1940 (Mrs.
Fry). Recorded in Panama August 17—April 28.

22 The yellowthroat (Geothlypis trichas) is mentioned in Chapman’s 1929
(but not in his 1938) list. As the only reported Panama specimen is a migrant
collected in western Chiriqui, the later omission may have been intentional.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 53

*Basileuterus delatrii mesochrysus: Chestnut-capped Warbler (Sclat-
er’s).

Collected June 18, July 27, 1927 (Van Tyne). According to Chap-
man, “not uncommon along clearing borders,” but I have not seen it
on any of my visits. It favors second-growth thickets and scrub.
Notes: Calls include a simple chit, a zeep-zeep or dzit-dzit; song is a
rather rapid, dry, chittering chit-cha-chup-cha-chuweép, with varia-
tions. Breeding: In Panama near the Canal Zone, juvenal being fed
June 25 (Eisenmann).

Family ICTERIDAE: Blackbirds, Troupials, and Allies

*Zarhynchus wagleri ridgwayi: Chestnut-headed Oropéndola (Wag-
ler’s).

Small bands are frequently seen around the clearing and in the for-
est ; location of breeding colonies not presently known. Noies: A slash-
ing, liquid croak, grwdckoo, grwackoo, grwak, grwak ; often grwdackoo
alone; also a series of gurgling notes like the sound of dripping water,
plup, plup, plup, plip-loo-vipoo; also plup alone. Breeding: In the
former colony near the laboratory nest-building regularly began the
first or second week of January, egg laying began in late January and
continued to April, and by the end of June the young had left (Chap-
man) ; June 27, 1925, a few young still in nest (Gross).

*Cacicus cela vitellinus: Yellow-rumped Cacique (Lawrence’s).

A local colonial nester, favoring trees with open surroundings or
towering above their neighbors. Breeding: A colony of 16 nests
attached to a bush growing on a dead trunk in water of the estero
west of Salud Point, April-May 1935, destroyed by a snake (Skutch).

Amblycercus holosericeus holosericeus: Yellow-billed Cacique (Prev-
ost’s).

Seen February 19, 1940, at edge of the clearing (Mrs. Fry). A com-
mon thicket bird in more open country. Notes: The male utters
mellow whistles, answered by the female with a long dry churr
(Skutch). Breeding: Not colonial, nest with eggs in the Canal Zone,
April 24 (Hallinan).

Psomocolax oryzivorus violeus: Giant Cowbird (Colombian Rice
Grackle).
Frequents breeding colonies of Zarhynchus and Cacicus, in whose
nests it lays its egg. After the breeding season it gathers in flocks,
feeding in fields and often attending cattle.

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS 1a7

Icterus galbula: Baltimore Oriole.

Winter visitant, February 11, 1929 (Chapman) ; February 8, 1940,
male and female (Mrs. Fry) ; March 24, 1950 (Cottrell). Recorded
in Panama October 2—April 20.

Icterus spurius: Orchard Oriole.

Winter visitant, February 19, 1937 (Chapman); February 22,
1948 (Wetmore) ; March 20, 1950, male and female (Cottrell). Re-
corded in Panama August 2—March 20.

Icterus mesomelas salvinii: Yellow-tailed Oriole (Salvin’s).

Infrequently noted near the laboratory. April 17, 1942 (Imhof) ;
March 12, 1946, March 8 and May 3, 1949 (Wetmore).

Icterus chrysater giraudiui: Yellow-backed Oriole (Giraud’s).

Common in and around the clearing and often along the lake shore.
Notes: Song is a rather hesitant, disconnected series of 6 to 14
loud, clear whistles, moving irregularly up and down scale, and end-
ing in an inconclusive manner. Another common call goes teéa, cheep-
cheep-cheep tee, the first and last notes whistled, the middle ones
nasal; often the last note is suppressed.

Family THRAUPIDAE: Tanagers

*Tanagra fulvicrissa: Fulvous-vented Euphonia.
Regular in small groups, feeding on mistletoe berries.

Tanagra lanurostris crassirostris: Thick-billed Euphonia.
“Not common” (Chapman). In the clearing, April 1935 (Skutch).

*Tangara inornata languens: Plain-colored Tanager.

Very common in small groups around the clearing. Notes: When
feeding, a low ist tst tst ; when flying off, tsrrr. Breeding: Nests found
in mango and lime trees near the laboratory, April and May 1935
(Skutch). In Canal Zone, fledgling being fed, August 17 (Har-
rower, Arbib).

Tangara lavinia lavinia: Rufous-winged Tanager (Lavinia’s).

Seen July 22, 1933 in the clearing (Arbib). A little-known forest
species.”

°3 The rather similar green tanager, 7. gyrola deleticia (not mentioned by

Sturgis), has also been recorded in the Canal Zone; it lacks the rufous wings
and has a blue rump.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 55

Tangara larvata franciscae: Golden-masked Tanager (Mrs. Wilson’s).

Rather common in small groups around the clearing. Breeding:
Nests found in orange and avocado trees near the laboratory, Feb-
ruary—May, 1937 (Skutch) ; collecting nesting material, July 13, 1947
(Mrs. Hobson).

*Thraupis episcopus diaconus: Blue-gray Tanager (Blue).

Very common in the clearing, and found in other open situations.
Notes: Song is a fast, sibilant, twittering tsu tsu tseéwee tsu-tseéwee
tsu-tseéwee tseéwee tseéwee, with variations. Calls are chup; also
chueép; also seeee. Breeding: Nests, January (Chapman), May to,
1935 (Skutch) ; building nest, July 4, 1948 (Longenecker, Eisen-
mann). In the Canal Zone, nests with eggs, April 13 and May 6
(Hallinan) ; in Panama City, feeding young in nest, June 23 (Eisen-
mann).

*Thraupis palmarum atripennis: Palm Tanager.

Very common in the clearing. Notes: Song somewhat like that
of the preceding species, but with a distinct “r’” sound: sésurt siisuri,
sree sree sree, susuri siisuri suisuri, with variations. Calls are see-ee
with a rising or questioning inflection, and see-you with a falling in-
flection. Breeding: In natural and artificial crannies, late in January
1926 (Chapman) ; February 28, 1935, in coco palm (Skutch) ; June 1,
1935, young hatched, nest in cranny in top of dead trunk (Skutch).

*Ramphocelus dimidiatus isthmicus: Crimson-backed Tanager.
Irregular, but sometimes fairly common in the clearing. A bird
of thickets and small trees in open country. Breeding: February and
April, 1935 ; second nest destroyed and same pair raised another brood
in May (Skutch). In the Canal Zone, nests with eggs found March 3
(Stone 1918), May to and 24 (Hallinan), June 11 (Stone, 1918).

Ramphocelus icteronotus: Yellow-rumped Tanager.

Irregular, but sometimes not uncommon around the edge of the
clearing. A bird of thickets at the edge of humid forest. Breeding:
In the Canal Zone, nests with eggs found April 30 and May 14
(Stone, 1918).

*Piranga rubra rubra: Summer Tanager.

Regular winter visitant, November 1939 (Skutch)—March 24, 1933
(Carleton), and March 24, 1950 (Cottrell). Recorded in Panama
September 3- April 16.

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

*Piranga olivacea: Scarlet Tanager.
Male collected April 5, 1936 (Niedrach). In Panama an uncom-
mon migrant recorded March 25—April 5.

*Habia fuscicauda erythrolaema: Dusky-tailed Ant-tanager.

Not uncommon in thickets in the forest. Notes: Song is a beautiful
flutelike warble. Call is a harsh wrenlike scold. Breeding: In the
Canal Zone, nest with eggs found May 14 (Stone, 1918). Some
authorities consider this form a race of the red-throated ant-tanager,
H. gutturalis.

Tachyphonus rufus: White-lined Tanager (Boddaert’s).

Irregular about the clearing ; a bird of thickets at the edge of wood-
land. Breeding: In the Canal Zone, nest with eggs found May 7
(Stone, 1918).

*Tachyphonus luctuosus panamensis: White-shouldered Tanager.

Not uncommon in the clearing and in lighter woods; a bird of
forest edge. Notes: tsip, tsip.

Heterospingus rubrifrons: Sulphur-rumped Tanager.

Seen (4 birds) November 20, 1939, in the tree-tops in the forest
near the tower (Skutch). A little-known species. Male: Above
sooty-slate with yellow rump; below slate-gray, tinged with yellow
on abdomen, with white patch on each side of breast; white under
wing coverts. Female: Duller and less yellow on abdomen. Length,
6 inches. Considered by Hellmayr a “hen-feathered” race of H.
avanthopygus.

*Eucometis penicillata cristata: Gray-headed Tanager (Gray-crested).

Fairly common in light woodland and occasional at the edge of the
clearing. A bird of shrubbery and low trees. Notes: chewp, chewp;
also a sharp chip. Breeding: March 29, 1935 (Skutch) ; July 1, 1948,
nest with one egg being incubated at Snyder-Molino trail O, hatched
July 11 (Longenecker, Eisenmann).

*Rhodinocichla rosea eximia: Rose-breasted Thrush-tanager (Pan-
ama).
One at border of clearing winter of 1936-1937; found dead Feb-
ruary 17, 1937 (Chapman, Weber). A bird of dense thickets, most
common on the Pacific slope. Notes: A rich, very loud, rather hollow,

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 57

whistled chd-ho, repeated over and over, and sometimes varied to
cheéa-wo, cho-wéy.

Family FRINGILLIDAE: Finches, Grosbeaks, and Allies

*Saltator maximus intermedius: Buff-throated Saltator.

Occasional in thickets around the clearing. Breeding: Carrying
nesting material April 15, 1935; nest with nestling in lime tree May
9, 1935 (Skutch). In the Canal Zone, nests with eggs found April 1
(Hallinan) and April 30 (Stone, 1918).

*Saltator albicollis isthmicus: Streaked Saltator.

Occasional in the thickets around the clearing; common in open
country. Notes: Aloud, whistled cheé-o0,cheé-oo, cheé-oo, cheéeee-oo;
or choo, cheé-oo, cheé-oo, cheéee-oo, varied at times to chip-cheé-up,
chip-cheé-up, chup-cheé-oo; also a sharp tseek. Breeding: In the
Canal Zone, nests with eggs found April 7, May 1o (Hallinan), Au-

gust 10 (Harrower).

*Pitylus grossus saturatus: Slate-colored Grosbeak.

Common in the forest, particularly near the clearing ; more often
heard than seen. Notes: A variety of high, whistled phrases, witchee-
weéoo, cheéoo; also wee-ee, witchoo cheéoo cheer; also hod-ee, weéoo-
wihoo-wéyoo. Call is a metallic chip.

*Cyanocompsa cyanoides cyanoides: Blue-black Grosbeak (Panama
Blue).

Common in thickets at the edge of the clearing and in the lighter
woodland. Notes: Song a series of loud, clear, deliberate whistles
do-do, deh, dee, deh, do, often followed by a soft, irregular twitter ;
sometimes there are only five whistled notes, corresponding very well
with the solmization “do do re mi re.” Also uttered is a sharp scold-
ing kee-eey. Breeding: Nest with eggs, July 1947 (C. A. Moore).
In the Canal Zone, nest with two fresh eggs, July 27, 1933 (Har-
rower).

Spiza americana: Dickcissel.

Seen twice in winter (Chapman). In Panama a winter visitant,
numerous locally on migration, September 11—May 10, one specimen
June 16.

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

*Sporophila aurita aurita: Variable Seedeater (Hicks’s).

Very common in the clearing. Notes: A sweet, rapid, twittering,
somewhat canarylike tsiwee tsiwee tsiwee, chee chee chee, twee-iwee-
twee-twee, chirr chirr chirr chirr with variations, the chirrs often
omitted. Call is a sweet questioning chiwee or tsiwee; also cheep.
Breeding: Nests with fresh eggs, June, July, August, September
(Gross, Eisenmann); young left nest as early as June 26, 1951
(Laughlin, Eisenmann), and there were young in the nests as late as
second week of October 1927 (Gross). In the Canal Zone, nests with
eggs found as early as last week of May, but eggs are laid chiefly
June-August (fide Gross).

Sporophila nigricollis nigricollis: Yellow-bellied Seedeater.

One seen in the clearing March 19, 1933 (Carleton). A common
open-country species, often noted at Frijoles. Notes: Song is sweet,
but ends usually with two buzzy phrases, seéwee-seéwee-seéwee-
seéwee soosoo, bzeéwee-bzeéwee; another song is zeéoo, zeéoo, bzii,

zu; a third is wit-seéoo, tsitseéoo, tsee. Breeding: In the Canal
Zone and Panama City, nests with eggs, July 13 (Harrower, Arbib),
July 17 (Eisenmann).

*V olatima jacarina splendens: Blue-black Grassquit.

Adult male collected April 27, 1927 (Van Tyne). On the main-
land an abundant bird of grassy areas. Notes: An explosive bzeé-eep
given by the male as it jumps a foot in the air, often repeating the
performance every 10 to 15 seconds for a half-hour or more. Breed-
ing: In the Canal Zone and nearby Panama, occupied nests, July 2
(eggs) (Stone, 1918) ; July 6 (2 eggs) hatched July 7, July 13 (3
young), July 14 (1 young and I egg) (Eisenmann).

Oryzoborus funereus: Thick-billed Seed-finch (Rice Grosbeak).

Occasional in the clearing. An open-country species. Notes: Song
is a sweet warble, given from the top of a small tree or other eleva-
tion. Breeding: In the Canal Zone, nest with eggs found April 11
(Hallinan).

Arremonops conirostris striaticeps: Green-backed Sparrow (Lafres-
naye’s).
Occasional in thickets of the clearing ; a bird of more open country.
Notes: Song, a loud, clear note, repeated at first slowly then gradually
faster and faster, cho, cho, cho, cho cho cho cho-cho-chochocho.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISEN MANN 59

Calls, whit-cho; also a whistled whit or white (like the white in the
bob-white call) ; also whup; also cheéoo, cheéoo; also chep-chep-chep.
Breeding: July 10, 1950, juvenal with adult (Eisenmann). In the
Canal Zone and Panama City, nests with eggs, April 18, June 2, June
11 (Stone, 1918), July 16 (Harrower, Arbib); nests with young,
June 20, July 17 (Eisenmann, I. R. Eisenmann, Jr).

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KeEnoyer, L. A.
1929. General and successional ecology of the lower tropical rain forest of
Barro Colorado Island, Panama. Ecology, vol. 10, No. 2.
LEHMANN, F. C.
1943. El genero Morphnus. Caldasia, vol. 7, pp. 165-170.
Moore, C. M.
1948. Choosing a bird for special study on Barro Colorado Island. Indiana
Audubon Soc. Year Book, 1948, pp. 24-26.
PETERS) Jae:
1931-1951. Check-list of birds of the world, vols. 1-7. Harvard Univ. Press,
Cambridge, Mass.

NO. 5 BIRDS OF BARRO COLORADO ISLAND—EISENMANN 61

QUAINTANCE, C. W.
1949. Vignettes of Barro Colorado life. Turtox News, vol. 27, pp. 74-78.
RipGway, Rosert.
1901-1950. Birds of North and Middle America. U. S. Nat. Mus. Bull. 50,
pts. I-11. (Continued by H. Friedmann.)
Rocers, C. H.
1939. The swifts of Panama. Auk, vol. 56, pp. 81-83.
SxutcH, A. F.
1931. Life history of Rieffer’s hummingbird. Auk, vol. 48, pp. 481-500.
1934. A nesting of the slaty antshrike on Barro Colorado Island. Auk,
vol. 51, p. 9.
1935. Helpers at the nest. Auk, vol. 52, pp. 252-273.
1940. Social and sleeping habits of Central American wrens. Auk, vol. 57,
Pp. 203-312.
1943. The family life of Central American woodpeckers. Sci. Month.,
vol. 56, pp. 358-364.
1945. On the habits and nest of the antthrush Formicarius analis. Wilson
Bull., vol. 57, pp. 122-128.
1946. Life histories of two Panamanian antbirds. Condor, vol. 48, pp. 16-28.
1949. Life history of the yellow-thighed manakin. Auk, vol. 66, pp. 1-24.
1950a. The nesting seasons of Central American birds in relation to climate
and food supply. Ibis (1950), pp. 185-222.
1950b. An adventure with toucans. Nature Mag., vol. 43, pp. 410-413, 440.
1951a. Congeneric birds nesting together in Central America. Condor, vol.
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1951b. Life history of the boat-billed flycatcher. Auk, vol. 68, pp. 30-49.
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1933. The flora of Barro Colorado Island, Panama. Contr. Arnold Arb.,
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1918. Birds of the Panama Canal Zone, with special reference to a collec-
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Stureis, B. B.
1928. Field book of birds of the Panama Canal Zone. New York.
Sutton, G. M.
1951. Mexican birds. Univ. Oklahoma Press, Norman, Okla.
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62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I1I7

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1948. Studies of Peruvian birds. No. 53. The family Trogonidae. Amer.
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1449, Pp. 19-29.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 6

fib SCAPHOPOD) MOLLUSKS COLLECTED BY
THE FIRST JOHNSON-SMITHSONIAN
DEEP-SEA EXPEDITION

(WitTH ONE PLATE)

BY
WILLIAM K. EMERSON

Allan Hancock Foundation
University of Southern California

(PusticaTion 4059)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
FEBRUARY 26, 1952

The Lord Waltimore Press

BALTIMORE, MD., U. S&S A.

Tie SCAPHORODI MOLLUSKS COLLECTED BY
THE FIRST JOHNSON-SMITHSONIAN
DEBE-SEN APE Dron:

By WILLIAM K. EMERSON

Allan Hancock Foundation
University of Southern California

(WitTH ONE PLATE)

A small but significant collection of scaphopods was obtained in
1933 by the First Johnson-Smithsonian Deep-Sea Expedition to the
Puerto Rican Deep, sponsored by the late Eldridge R. Johnson, of
Philadelphia. Although only one new species was found, examples
were obtained of several species that had been represented previously
by unique specimens or by very few individuals. A better taxonomic
understanding of several formerly little-known species has been made
possible owing to the acquisition of numerous examples of these spe-
cies. The expedition procured a total of 17 species, 2 of which are
questionably identified. The species are listed both systematically and
by station. The station data shed little light upon the natural associa-
tions of the species because of the great vertical depth range encom-
passed by many of the hauls. The specimens are all in the collection
of the division of mollusks of the United States National Museum.

No attempt has been made to compile a complete synonymy of the
species. The reader is referred to the excellent monograph of the
eastern American scaphopods by John B. Henderson.? Reference is
made to the original descriptions and to Henderson for the complete
synonymies and descriptions. The classification used is essentially
that of Henderson.

The author wishes to express his appreciation to Dr. Harald A.
Rehder, curator, division of mollusks, United States National Mu-
seum, for having extended to him the use of the laboratory facilities

1 Contribution No. 62 of the Allan Hancock Foundation, University of South-
ern California. Previous papers on the scientific results of this expedition were
published in Smithsonian Misc. Coll., vol. 91. This report on the scaphopod
mollusks was not completed in time to be included in that volume.

2 Henderson, John B., A monograph of the east American scaphopod mollusks,
U. S. Nat. Mus. Bull. 111, pp. i-vi, 1-177, pls. 1-20, 1920.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 6

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

and collections in making this study, and to the administration of the
Allan Hancock Foundation for material aid in its completion.

Family DENTALIIDAE
Genus DENTALIUM Linné, 1758

Dentalium LinnE, Syst. Nat., ed. 10, p. 785, 1758.
Dentalium MontrFort, Conchyl. Syst., vol. 2, p. 23, 1810.

Genotype by subsequent designation, Montfort 1810, Dentalium ele-
phantium Linné, 1758; Recent, Amboina and Philippine Islands.

DENTALIUM (DENTALIUM) GOULDII PORTORICENSE Henderson, 1920

Dentalium (Dentalium) gouldti portoricense HENpERSON, U. S. Nat. Mus. Bull.
III, p. 30, pl. 2, fig. 5, 1920 (Mayagiiez Harbor, Puerto Rico).

Remarks.—A large series of over 100 specimens of this rare shell
was taken at one station. Only five individuals were known previously.
The material lacks the well-developed intercostal longitudinal striae
of the typical subspecies. The hexagonal tip is simple, there being
no slit or notch. Fresh specimens have a vitreous shell that has a
glazed appearance.

Records.—Station 10, 18°209'20” N., 66°05/30” W., 120-160 fath-
oms (4). Station 26, 18°30’20” N., 66°22'05” W., 33-40 fathoms (1).
Station 104, 18°30'40” N., 66°13'20” W., 80-120 fathoms (100+).

Subgenus COCCODENTALIUM Sacco, 1896

Coccodentalium Sacco, Boll. Mus. Univ. Torino, vol. 11, p. 98, 1896.
Coccodentalium Sacco, Moll. Terr. Terz. Piedmonte e della Liguria, pt. 22,
p. I10, 1897.

Subgenotype by original designation, Dentalium radula Schroeter,
1784; Miocene, Piedmont of Italy.

DENTALIUM (COCCODENTALIUM) CARDUUS Dall, 1889

Dentalium carduus Dati, Bull. Mus. Comp. Zool., vol. 18, pt. 2, p. 423, pl. 27,
fig. 3, 1889 (off Santa Lucia).

Dentalium (Dentaliwm) carduus, HENDERSON, U. S. Nat. Mus. Bull. 111, p. 33,
pl. 3, figs. 4, 5, 7, 1920 (Little Bahama Bank; off Grenada).

Remarks.—A fragment representing the anterior portion of this
rare species was collected. This specimen appears to be the sixth
record, as Henderson (1920) states the species to be known from only
five examples. This species is here placed in the subgenus Coccoden-

NO. 6 SCAPHOPOD MOLLUSKS—EMERSON 3

taliuvm because of the possession of surface sculpture similar to Den-
talium radula Schroeter, the subgenotype.

Record.—Station 100, 18°38’45” N., 64°53'45” W., 100-300 fath-
oms (I).

Subgenus DENTALE Da Costa, 1778 3

Dentale Da Costa, Hist. Nat. Test. Brit., p. 24, 1778.
Antalis HERRMANNSEN, Indicis Generum Malacoz., vol. 1, p. 63, 1846.
Antalis Pitspry and SHarp, Man. Conch., vol. 17, p. 37, 1897.

Subgenotype by monotypy, Dentale vulgare Da Costa 1778; Re-
cent, Mediterranean and Adriatic Seas, Atlantic Ocean from Spain to
Belgium, etc.; Tertiary of Belgium and Italy.

DENTALIUM (DENTALE) BARTLETTI Henderson, 1920

Dentalium (Antalis) bartletti HENpErRson, U. S. Nat. Mus. Bull. 111, p. 55,

pl. 8, figs. 2, 7, 1920.

Remarks.—Only one lot containing five examples of this fragile
deep-water species was taken. Henderson’s (1920) West Indian rec-
ords include specimens from off Havana, St. Vincent, and Martinique,
in 357 to 464 fathoms.

Record.—Station 93, 18°38’00” N., 65°09'30” W., 350-400 fath-
oms, “cement-like mud” (5).

DENTALIUM (DENTALE) CERATUM Dall, 1881
Dentaliwm ceratum Dat, Bull. Mus. Comp. Zool., vol. 9, p. 38, 1881.
Dentalium (Antalis) ceratwm, HENDERSON, U. S. Nat. Mus. Bull. 111, p. 40,

Dl 7a hes. 2) As 5, O07. TO20:

Remarks.—Henderson divided this extremely variable species into
several subspecies of doubtful validity. Though the few specimens
obtained are poorly preserved, they appear to represent two of Hen-
derson’s subspecies, his “typical” and his southern “geographical race”
from Barbados, Dentalium ceratum tenax.

Records.—Station 102, 18°50'30” N., 64°43'00” W., 90-500 fath-
oms (4). Station 104, 18°30’40” N., 66°13'20” W., 80-120 fathoms
(2).

Subgenus LAEVIDENTALIUM Cossmann, 1888
Laevidentalium CossMANN, Ann. Soc. Roy. Malacol. Belgique, vol. 23, p. 7,

1888.

Laevidentalium .HeNnprrson, U. S. Nat. Mus. Bull. 111, p. 73, 1920.

Subgenotype by original designation, Dentalium incertum Deshayes,
1825 ; Eocene of the Paris Basin.

8 For the extensive synonymy of Dentale, see Emerson, Nautilus, vol. 64,
Pal7, TO5T.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

DENTALIUM (LAEVIDENTALIUM) CALLIPEPLUM Dall, 1889

Dentalium callipeplum Dati, Bull. Mus. Comp. Zool., vol. 18, p. 419, pl. 27,
fig. 12(b), 1880.

Dentalium (Laevidentalium) callipeplum, HENpERSoN, U. S. Nat. Mus. Bull.
TDL, p. 74, ple 12) ehiee 5.) 1920:

Remarks.—A good series of well-preserved specimens representing
this rather uncommon species was taken. The specimens range in size
from fragments of the small, narrow, needlelike anterior portions to
large, entire individuals, one of which is apparently the longest speci-
men known, measuring nearly 75 mm. in length. The specimens in a
good state of preservation show clearly the development of microscopic
incremental rings. These rings are in close association posteriorly and
become progressively farther apart anteriorly for approximately one-
third the length of the shell, after which they become irregularly
spaced or nearly completely obscure in the remaining portion. The
annulations are produced by slightly oblique incised lines. In adult
specimens, the posterior orifice generally has a shallow, subtriangular
notch on the concave face.

Records.—Station 23, 18°32’15” N., 66°17’45” W., 260-360 fath-
oms (8). Station 25, 18°32'15” N., 66°22’10” W., 240-300 fathoms
(4). Station 62, 19°25’45” N., 69°09’00” W., 350 fathoms (10).
Station 94, 18°37'45” N., 65°05’00” W., 300-470 fathoms (1).

DENTALIUM (LAEVIDENTALIUM) ?PERLONGUM Dall, 1881

Dentalium perlongum Dati, Bull. Mus. Comp. Zool., vol. 5, No. 6, p. 61, 1878
[name only].

Dentalium perlongum Datu, Bull. Mus. Comp. Zool., vol. 9, p. 36, 1881.

Dentalium (Laevidentalium) perlongum, HENbDERSON, U. S. Nat. Mus. Bull. 111,

p. 75, pl. 9, fig. 1, 1920.

Remarks.—A few small, glossy, needlelike fragments which may
represent the posterior tips of this species were taken. The largest
fragment measures only 12 mm. in length.

Record.—Station 13, 18°31'05” N., 66°02'15” W., 200-300 fath-
oms (7).

Subgenus EPISIPHON Pilsbry and Sharp, 1897

Episiphon Pirspry and SHarp, Man. Conch., vol. 17, p. 117, 1897.
Episiphon Suter, Man. New Zealand Moll., p. 821, 1913.

Subgenotype by subsequent designation, Suter, 1913, Dentalium
sowerbyi Guilding, 1834; Recent, southeastern United States and
West Indies.

NO. 6 SCAPHOPOD MOLLUSKS—EMERSON 5

DENTALIUM (EPISIPHON) JOHNSONI Emerson, new species
Pity, fips

Diagnosis.—Shell slender, thin but not fragile, slightly but uni-
formly curved, strongly compressed laterally to form an ovate section ;
section more ovate posteriorly than anteriorly. Juvenile specimens
slowly and regularly increasing in diameter, apex attenuated to a
needlelike orifice, generally with no indication of a pipe; apex in adult
stage often truncated with a very thin, rounded inner tube situated
slightly off center toward the convex face and projecting from the
orifice. Apical portion of mature individuals filled except for the space
occupied by the tube. Aperture of truncated specimens only slightly
larger in diameter than the apical orifice. Shell white, vitreous, semi-
transparent or translucent where clouded by semiopaque growth rings.
Surface glossy, essentially smooth, broken only by occasional irregular
growth rings, without longitudinal sculpture. Measurements of the
holotype: length, 24 mm.; diameter of apical orifice, 1 mm.; diameter
of aperture, 1.3 mm. Mature specimens range from 17 to 28 mm. in
length.

Remarks.—This relatively large species is one of the largest mem-
bers of a subgenus composed primarily of small forms. However, it
meets the other requirements of the group. The degree of curvature
varies slightly with individuals, a few being moderately curved.

Comparisons.—No representative described from the Atlantic Ocean
reaches the large size of this species. Dentaliwm (Episiphon) sower-
byi,* from off the Florida coast and in the West Indies, is a minute
species characterized by prominent, coarsely developed growth rings
and a circular outline. The most closely related form among the fossil
species of the Caribbean region appears to be the Miocene species,
Dentalium (Episiphon) macilentum,® which is a much smaller, more
compressed species.

Type locality —Off Puerto Rico, Station 25, 18°32’15” N., 66°22’
10” W., 240-300 fathoms.

Type depository—Holotype, U.S.N.M. No. 603543, Station 25,
18°32'15”. N., 66°22’10” W., 240-300 fathoms. Paratypes, U.S.N.M.
No. 429714, Station 25, 18°32’15” N., 66°22'10” W., 240-300 fathoms
(72).

Other records.—Station 12, 18°31'00” N., 66°00’15” W., 200-300
fathoms, blue mud (1). Station 13, 18°31’05” N., 66°02’15” W.,

4 Guilding, Trans. Linn. Soc. London, vol. 17, p. 35, pl. 3, fig. 7, 1834.
5 Pilsbry, Proc. Acad. Nat. Sci. Philadelphia, vol. 63, pp. 166-167, fig. 1-2,
I911 (Bowden, Jamaica).

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

200-300 fathoms, blue mud (12). Station 14, 18°31’00” N., 66°04’10”
W., 240-340 fathoms (3). Station 23, 18°32’15” N., 66°17’45” W.,
260-360 fathoms (4). Station 32, 18°25’50” N., 67°14'55” W., 200-
280 fathoms (1). Station 67, 18°30'12” N., 65°45'48” W., 180-280
fathoms, mud (3). Station 84, 18°32’30” N., 65°18’30” W., 300-350
fathoms (1). Station 93, 18°38’00” N., 65°09'30” W., 350-400 fath-
oms, mud (30). Station 94, 18°37’45” N., 65°05’30” W., 300-470
fathoms (1). '

Subgenus BATHOXIPHUS Pilsbry and Sharp, 1897

Bathoxiphus Prrspry and SHARP, Man. Conch., vol. 17, p. 121, 1897.
Bathoxiphus Botssevain, Siboga Exped., vol. 54, Scaphopoda, p. 48, 1906.

Subgenotype by subsequent designation, Boissevain 1906, Den-
talium ensiculus Jeffreys, 1877; Recent, Atlantic Ocean, in deep water.

DENTALIUM (BATHOXIPHUS) ENSICULUS Jeffreys, 1877

Dentalium ensiculus JEFFREYS, Ann. Mag. Nat. Hist., ser. 4, vol. 10, p. 154,
1877.
Dentalium (Bathoxiphus) ensiculus, HENDERSON, U. S. Nat. Mus. Bull. 111,

p. 81, pl. 14, figs. 1, 4, 5, 7, 9, 1920.

Remarks.—One lot containing 50 specimens was dredged in 350
to 400 fathoms. The character of these specimens substantiates Hen-
derson’s findings that the Antillean material tends to be somewhat
more slender than the northern representatives. The largest specimen
measures 30 mm. in length. This deep-water species apparently has
an extensive geographical range, having been previously reported
from off the New England coast to St. Bartholomew, W. I., and in
the eastern Atlantic off the coasts of Portugal, Ireland, etc.

Record.—Station 93, 18°38’00” N., 65°09’30” W., 350-400 fath-
oms, “cement-like mud” (50).

Subgenus COMPRESSIDENS Pilsbry and Sharp, 1897
Compressidens Pitspry and SHarp, Man. Conch., vol. 17, p. 123, 1897.

Type by original designation, Dentalium pressum Pilsbry and
Sharp, 1897; Recent, West Indies to the Florida Keys region.

Remarks.—The western Atlantic representatives of this group may
actually be members of the Siphonodentaliidae of the subgenus Pul-
sellum. However, since the soft parts are not available for study, the
classification of Henderson (1920) is being followed.

SMITHSONIAN MISCELLANEOUS COLLECTIONS MO at’, IOs yy Vee al

SCAPHOPOD MOLLUSKS

I, Specimens of Entalina platamodes (Watson) shown attached to the mem-
branous tubes of a polychaetous worm. 2, A series of Dentalium (Episiphon)
johnsoni, new species, showing various stages of development and individual
variation; specimen at extreme right is the holotype, the remaining specimens
are paratypes.

NO. 6 SCAPHOPOD MOLLUSKS—EMERSON 7

DENTALIUM (COMPRESSIDENS) PRESSUM Pilsbry and Sharp, 1897

Dentalium compressum Watson, Journ. Linn. Soc. Zool. London, vol. 14, p.
516, 1879 [not Orbigny, 1850].

Dentalium (Compressidens) pressum PILtspry and SHARP, Man. Conch., vol. 17,
p. 124, pl. 7, fig. 11; pl. 22, figs. 50-52, 1897.

Dentalium (Compressidens) pressum, HENDERSON, U. S. Nat. Mus. Bull. 111,
p. 83, pl. 14, figs. 3, 6, 8, 1920.

RRemarks.—Only one small specimen of this Antillean-continental
slope species was found. It ranges north within the Gulf Stream to
the Florida Keys.

Record.—Station 67, 18°30'12” N., 65°45’48” W., 180-280 fath-
oms, mud (1).

DENTALIUM (COMPRESSIDENS) OPHIODON Dall, 1881

Dentalium ophiodon Dat, Bull. Mus. Comp. Zool., vol. 9, p. 38, 1881.
Dentalium (Compressidens) ophiodon, HENvERSON, U. S. Nat. Mus. Bull. 111,
p. 84, pl. 14, fig. 2, 1920.

Remarks.—A representative series of this Antillean-continental
slope species was taken. It is a smaller, more slender species with a
much less degree of compression than Dentalium (Compressidens)
pressum Pilsbry and Sharp, which has a similar range.

Records.—Station 13, 18°31'05” N., 66°02'15” W., 200-300 fath-
oms, “blue mud” (5). Station 25, 18°32'15” N., 66°22’10” W., 240-
300 fathoms, “‘soft mud” (6).

Family SIPHONODENTALIIDAE
Genus ENTALINA Monterosato, 1872

Entalina MontTerosato, Notizie intorno alle Conch. Fossile di Monte Pellegrino
e Ficarazzi, p. 27, 1872.

Entalina Sacco, Moll. Terr. Terz. Piedmonte e della Liguria, pt. 22, p. 114,
1807.

Genotype by subsequent designation, Sacco, 1897, Dentalium tet-
vagona Brocchi, 1814 (= ?Dentalium quinquangulare Forbes, 1843) ;
Miocene, northern Italy.

ENTALINA PLATAMODES (Watson), 1879

Siphodentalium [sic] platamodes Watson, Journ. Linn. Soc. Zool. London, vol.
14, DP. 510, 1879.

Entalina platamodes, HENDERSON, U. S. Nat. Mus. Bull. 111, p. 87, pl. 15, figs.
Bed i5; 7; 1020.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Entalina quadrata HEeNpverson, U. S. Nat. Mus. Bull. 111, p. 88, pl. 15, figs.
293); o10;9 1020:

Remarks.—Five lots totaling over 200 specimens of this unusual
species were taken. A number of dead specimens were found attached
to the sides of membranous tubes apparently belonging to a poly-
chaetous annelid of the genus Nothria ° (see pl. 1, fig. 1). This appears
to be the first record of such an association.

Watson (1879) described Entalina platamodes (off Culebra Island,
West Indies), from a poorly preserved specimen as indicated by his
remarks, “neither end is fresh enough for description.” However, the
general characters of the shell were clearly defined by his statement,
“shell five sided, with four sharp corners, which are nearly right
angles, and one very obtuse angle along the concave curve . . . there
are a few longitudinal striae . . . strongest near the angles, more or
less obsolete as they recede from these.” Henderson (1920) described
Entalina quadrata from one well-preserved specimen which possesses
a continuous quadrate section and many longitudinal riblets. The
material at hand shows that the concave face varies considerably in
the degree of medial keel development. Most specimens appear to be
five-sided, owing to the presence of this ridge. However, some appear
to have four sides when the ridge is not well developed. While the
arrangement and number of the longitudinal riblets is more constant in
most specimens, the pattern is often interrupted by wear. Apparently
Watson’s type specimens were badly worn, with the result that the
riblets were less discernible, while Henderson’s type specimen of E.
quadrata chanced to be a well-preserved individual showing the com-
plete arrangement of the riblets. Since the intergrading specimens
appear to connect the two forms, the species are here considered to be
conspecific.

This species is closely related to Entalina quinquangularis (Forbes)
from the Mediterranean-Aegean Seas, and the northwestern Atlantic
Ocean.

Records.—Station 13, 18°31’05” N., 66°02’15” W., 200-300 fath-
oms, blue mud (25). Station 23, 18°32'15” N., 66°17’45” W., 260-
360 fathoms, mud (40+). Station 25, 18°32’15” N., 66°22’10” W.,
240-300 fathoms, mud (60). Station 35, 18°23'40” N., 67°16'45” W.,
180-280 fathoms (3). Station 67, 18°30’12” N., 65°45/48” W., 180-
280 fathoms, mud (100+).

6 Provisionally identified by Dr. Olga Hartman, Allan Hancock Foundation,
University of Southern California.

No. 6 SCAPHOPOD MOLLUSKS—EMERSON 9

Genus CADULUS Philippi, 1844
Cadulus Puitipp1, Enumeratio Molluscorum Sicilae, vol. 2, p. 209, 1844.

Genotype by monotypy, Dentalium ovulum Philippi, 1844 (=Cad-
ulus (Cadulus) ovulum Philippi) ; Recent, Mediterranean Sea.

CADULUS (CADULUS) CONGRUENS Watson, 1879

Cadulus curtus congruens Watson, Journ. Linn. Soc. Zool. London, vol. 14,

Pp. 527, 1879.

Cadulus (Cadulus) congruens, Henverson, U. S. Nat. Mus. Bull. 111, p. 142

pl. 20, fig. 10, 1920.

Remarks.—One lot containing 18 specimens, which apparently
represent this species, was dredged in fairly deep water. This species
was not previously represented in the United States National Museum
collection.

The types were collected by the Challenger off Culebra Island, West
Indies, in 390 fathoms with a pteropod-00ze bottom.

Record.—Station 67, 18°30'12” N., 65°45'48” W., 180-280 fath-
oms, mud (18).

CADULUS (CADULUS) EXIGUUS Watson, 1879

Cadulus exiguus Watson, Journ. Linn. Soc. Zool. London, vol. 14, p. 528, 1870.
Cadulus (Cadulus) exiguus, HENDERSON, U. S. Nat. Mus. Bull. 111, p. 145, pl. 20,

fig. 9, 1920.

Remarks.—Three lots totaling 32 specimens were taken. This
species was previously represented by only two specimens in the
United States National Museum collection.

The type locality is off Culebra Island, West Indies, in 390 fathoms.

Records.—Station 13, 18°31’05” N., 66°02’15” W., 200-300 fath-
oms, blue mud (4). Station 35, 18°23’40” N., 67°16'45” W., 180-
280 fathoms (5). Station 67, 18°30’12” N., 65°45’48” W., 180-280
fathoms (23).

CADULUS (CADULUS) ?TERSUS Henderson, 1920

Cadulus (Cadulus) tersus HENpERsON, U. S. Nat. Mus. Bull. 111, p. 149, pl. 20,
fig. 4, 1920 (Barbados).

Remarks——One fragment which may represent this species was
dredged.

Record.—Station 67, 18°30'12” N., 65°45'48” W., 180-280 fathoms,
mud (1).

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Subgenus GADILOPSIS Woodring, 1925
Gadilopsis Woopr1NnG, Carnegie Inst. Washington Publ. 366, p. 206, 1925.

Subgenotype by original designation, Ditrupa dentalina Guppy,
1873. (=Cadulus (Gadilopsis) dentalinus (Guppy)); Miocene,
Jamaica.

CADULUS (GADILOPSIS) ACUS Dall, 1889

Cadulus acus DALL, Bull. Mus. Comp. Zool., vol. 18, p. 432, pl. 27, fig. 11, 1880.
Cadulus (Gadila) acus, HENvDERSON, U. S. Nat. Mus. Bull. 111, p. 140, pl. 20,
figs. II, 13, 1920.

Remarks.—Five lots of this uncommon species were taken. In this
shallow-water species the oblique growth rings are developed moder-
ately. There is a slight variation in width among individuals.

Records—All Samana Bay, Dominican Republic. Station 55, 19°
10'12” N., 69°27'/03” W., 17 fathoms (15). Station 56, 19°10'15”
N.,. 69°27'20” W057 fathoms | (352:). Station57,/ 1920207
69° 28/35” W., 18 fathoms, mud (2). Station 58, 19°10’20” N., 69°
29/15” W., 18 fathoms, mud (4). Station 59, 19°10'25” N., 69°30'05”
W., 18-19 fathoms, mud (2).

Subgenus PLATYSCHIDES Henderson, 1920
Platyschides HENvERsoN, U. S. Nat. Mus. Bull. 111, p. 104, 1920.

Subgenotype by original designation, Cadulus grandis Verrill, 1884;
Recent, west Atlantic, north of Hatteras.

CADULUS (PLATYSCHIDES) ELEPHAS Henderson, 1920

Cadulus (Platyschides) elephas HENvERsoN, U. S. Nat. Mus. Bull. 111, p. 107,
pl. 17, fig. 10, 1920.

Remarks.—Six specimens were obtained that can be referred un-
questionably to this species. It was previously known only from the
unique, which was dredged in 464 fathoms off St. Vincent.

This large ivory-white shell appears to have a simple posterior
orifice. The largest specimen measures 16 mm. in length, 1.5 mm.
less than the type. In these additional specimens the hemisphere on
the concave face is slightly more bulbous than in the type.

Records.—Station 25, 18°32'15” N., 66°22’10” W., 240-300 fath-
oms, mud (4). Station 67, 18°30'12” N., 65°45’48” W., 180-280
fathoms, mud (2).

NO. 6 SCAPHOPOD MOLLUSKS—-EMERSON II

CADULUS (PLATYSCHIDES) ?BUSHII Dall, 1889

Cadulus carolinensis bushii Dat, Bull. Mus. Comp. Zool., vol. 17, p. 430, 1889
(Barbados).

Cadulus (Platyschides) bushti, HENDERSON, U. S. Nat. Mus. Bull. 111, p. 1235,
pl. 19, fig. 10, 1920.

Remarks.—tThe one lot collected is questionably referred to this
species. While the specimens are about the same size and have an
outline similar to Cadulus bushi, they are heavy, rather opaque shells
in which the oral aperture is quite constricted.

This may prove to be a new species.

Record.—Station 21, 18°30'20” N., 66°10’30” W., 140-200 fath-
oms (8).

LIST OF SPECIES COLLECTED, ARRANGED BY STATIONS?

Station 10. Lat. 18°29’20” N. Long. 66°05’30” W. February 2, 1933
Lat. 18°30’24” N. Long. 66°04’15” W.
Off San Juan, Puerto Rico, in 120-160 fathoms, tangle:
Dentalium (D.) gouldii portoricense.

Station 12. Lat. 18°31’00” N. Long. 66°00’15” W. February 2, 1933
Lat. '18°30’30” N. Long. 66°o1’45” W.
Off Punta Maldonado, Puerto Rico, in 200-300 fathoms, beam trawl,
blue mud:

Dentalium (Episiphon) johnsoni.

Station 13. Lat. 18°31'05” N. Long. 66°02'15” W. February 2, 1933
Lat. 18°30’30” N. Long. 66°04’05” W.
Off Punta Maldonado, Puerto Rico, in 200-300 fathoms, beam trawl,
blue mud:
Dentalium (Laevidentalium) ?perlongum.,
Dentalium (Episiphon) johnsoni.
Dentalium (Compressidens) ophiodon.
Entalina (E.) platamodes.
Cadulus (C.) exiguus.

Station 14. Lat. 18°31’00” N. Long. 66°04'10” W. February 2, 1933
Lat. 18°30’30” N. Long. 66°03'15” W.
Off Punta Maldonado, Puerto Rico, in 240-340 fathoms, beam trawl:
Dentalium (Episiphon) johnsoni.

Station 21. Lat. 18°30'20” N. Long. 66°10’30” W. February 4, 1933
Lat. 18°31’15” N. Long. 66°12’20” W.
Off Putna Salinas, Puerto Rico, in 140-200 fathoms, beam trawl:
Cadulus (Platyschides) ?bushii.

7 Bartsch, Paul, Station Records of the First Johnson-Smithsonian Deep-Sea
Expedition, Smithsonian Misc. Coll., vol. 91, No. 1, pp. I-31, I map, 1033.

Station 23.

Station 25.

Station 26.

Station 32.

Station 35.

Station 55.

Station 56.

Station 57.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Lat. 18°32'15” N. Long. 66°17'45” W. February 4, 1933
[at. 19732/007N- one. Go;2tns.) Wis
Off Punta Cerro Gordo, Puerto Rico, in 260-350 fathoms, otter
trawl:
Dentalium (Laevidentalium) callipeplum.
Dentalium (Episiphon) johnsoni.
Entalina (E.) platamodes.

ater s2 15). IN eleonei Gor e2urOr Ns February 7, 1933
Lat. 18°32'05” N. Long. 66°22'10” W.
Off Punta Cerro Gordo, Puerto Rico, in 240-300 fathoms, dredge,
mud:

Dentalium (Laevidentalium) callipeplum.

Dentalium (Episiphon) johnsoni.

Dentalium (Compressidens) ophiodon.

Entalina (E.) platamodes.

Cadulus (Platyschides) elephas.

Wat. 11623020 aNe elbones(60222,05)4 We February 7, 1933

Eat. 18°30.30" N.. Long: 660°23'05° W.

Off Punta Cerro Gordo, Puerto Rico, in 33-40 fathoms, dredge,
rock, etc.:

Dentalium (D.) gouldii portoricense.

MatiemSe25(50 Ne leones O7o Tash Ve February 9, 1933

Eatynos23'504 Na Wones O7517-3540 Wie

Off Punta Jiguero, Puerto Rico, in 200-280 fathoms, dredge:
Dentalium (Episiphon) johnsoni.

Lat. 18°23'40” N. Long. 67°16'45” W. February 9, 1933
Lat. 18°24'45” N. Long. 67°14'15° W.
Off Punta Jiguero, Puerto Rico, in 180-280 fathoms, beam trawl:
Entalina (E.) platamodes.
Cadulus (C.) exiguus.

Lat. 19°10'12” IN. Long. 67°27'03” W. February 16, 1933

Lat. 19°10'15” N. Long. 69°27'10” W.

Samana Bay, Dominican Republic, in 17 fathoms, dredge:
Cadulus (Gadilopsis) acus.

Lat. 19°10'15” N. Long. 69°27’20” W. February 16, 1933
Lat. 19°10'15” N. Long. 69°28’05” W.
Samana Bay, Dominican Republic, in 17 fathoms, dredge:

Cadulus (Gadilopsis) acus.

Lat. 19°10'20” N. Long. 69°28'35” W. February 16, 1933

Lat. 19°10'20” N. Long. 69°29'00” W.

Samana Bay, Dominican Republic, in 18 fathoms, dredge, mud:
Cadulus (Gadilopsis) acus.

NO. 6

Station 58

Station 59

Station 62

Station 67

SCAPHOPOD MOLLUSKS—EMERSON 13

. Lat. 19°10’20” N. Long. 69°29'15” W. February 16, 1933
Lat. 19°10'25” N. Long. 69°30'05” W.
Samana Bay, Dominican Republic, in 18 fathoms, dredge, mud:
Cadulus (Gadilopsis) acus.

. Lat. 19°10’25” N. Long. 609°30'05” W. February 16, 1933
Lat. 19°10'35” N. Long. 69°30'40” W.
Samana Bay, Dominican Republic, in 18-19 fathoms, dredge, mud:
Cadulus (Gadilopsis) acus.

. Lat. 19°25’45” N. Long. 69°09’00” W. February 18, 1933
Lat. 19°27'45” N. Long. 69°14’45” W.
Off Punta Pescadores, Dominican Republic, in 350 fathoms, dredge:
Dentalium (Laevidentalium) callipeplum.

. Lat. 18°30'12” N. Long. 65°45'48” W. February 23, 1933
aterSes2a8ue Nr leone 05704612, . We
Off Punta Picua, Puerto Rico, in 180-280 fathoms, dredge, mud:
Dentalium (Episiphon) johnsoni.
Dentalium (Compressidens) pressum.
Entalina (E.) platamodes.
Cadulus (C.) congruens.
Cadulus (C.) exiguus.
Cadulus (C.) ?tersus.
Cadulus (Platyschides) elephas.

Station 84. Lat. 18°32’30” N. Long. 65°18’30” W. February 26, 1933

Lat. 18°39’00” N. Long. 65°17’00” W.
North of Culebra Island, in 300-350 fathoms, otter trawl:
Dentalium (Episiphon) johnsoni.

Station 93. Lat. 18°38’00” N. Long. 65°09’30” W. March 2, 1933

Lat. 18°37’45” N. Long. 65°05’00” W.
North of St. Thomas, Virgin Islands, in 350-400 fathoms, dredge,
mud:
Dentalium (Dentale) bartletti.
Dentalium (Episiphon) johnsoni.
Dentalium (Bathoxiphus) ensiculus.

Station 94. Lat. 18°37’45” N. Long. 65°05’00” W. March 2, 1933

Lat. 18°39’00” N. Long. 65°03’30” W.

North of St. Thomas, Virgin Islands, in 300-470 fathoms, dredge:
Dentalium (Laevidentalium) callipeplum.
Dentalium (Episiphon) johnsoni.

Station 100. Lat. 18°38’45” N. Long. 64°52’45” W. March 4, 1033

Lat. 18°40’15” N. Long. 64°50'15” W.
North of St. Thomas, Virgin Islands, in 100-300 fathoms, otter
trawl:
Dentalium (Coccodentalium) carduus.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Station 102. Lat. 18°50’30” N. Long. 64°43'00” W. March 4, 1933
Lat. 18°51'00” N. Long. 64°33’00” W.
Northwest of Anegada Island, in 90-500 fathoms, otter trawl:
Dentalium (Dentale) ceratwm.

Station 104. Lat. 18°30'40” N. Long. 66°13/20” W. March 8, 1033
Lat. 18°30’10” N. Long. 66°13'50” W.
Off Punta Boca Juana, Puerto Rico, in 80-120 fathoms, oyster
dredge, mud:
Dentalium (D.) gouldti portoricense.
Dentalium (Dentale) ceratum.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 7

fOsT RELATIONSHIPS OF MOTHS
oe THE GENERA..DEPRESSARIA AND
SCONOPTERIX. WITH.DESCRIPTIONS
OP INEW SPECIES

BY
j.. Ps GATES ‘CLARKE

U. S. Bureau of Entomology and Plant Quarantine

(Pustication 4083)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
APRIL, 23; 1952

The Lord Baltimore Press

: BALTIMORE, MD., U. S. Ae

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HOST RELATIONSHIPS OF MOTHS GF TEE
GENERA DEPRESSARIA AND AGONOP-
TERI NV Lin DESCRIPTIONS OP
NEW SPECIES

BY jer GAMES 'CLARIKE
U. S. Bureau of Entomology and Plant Quarantine

(WitTH Srx PriatEs)

In 1932 I began collecting larvae of the species of Depressaria
and Agonopterix (Lepidoptera: Oecophoridae) for the purpose of
rearing the moths and gaining some knowledge of their habits and
distribution and with the hope that unknown species might be dis-
covered. It soon became evident that undescribed species of these
genera were numerous and that many of their habits were both
intriguing and informative. Although interrupted, sometimes for
several years, these investigations have continued sporadically over
the past two decades with moderate success.

During the summer of 1950, by means of a grant-in-aid made by
the American Philosophical Society which defrayed some of the
expense of an extended trip through several western States, I was
able to resume my explorations in this field.1_ My purpose, essentially,
was to determine if possible whether the long-suspected host speci-
ficity of many species of Depressaria, particularly those attached to
plants of the genus Cicuta, really existed. Unfortunately, only a
few species of this genus of plants were found, and so the project
was enlarged to include species of moths attached to other umbellif-
erous plants.

In the course of the summer’s field work plants were examined at
80 localities, as shown in the list beginning on page 5. Infestations

1] wish to express my giatitude to the American Philosophical Society for
the grant-in-aid that made these investigations possible and to the Smithsonian
Institution for administering the grant. Also, I have many times been indebted
to my friend Dr. Lincoln Constance, Department of Botany, University of
California, for determinations of plants and for his stimulating company in the
field. During the summer of 1950, in connection with the present studies, I
again received his generous help, and again I thank him. I am indebted to
Harry F. Clements for furnishing the photograph for plate 5, figure 2; all
other photographs and drawings herein are of my own making. Unless other-
wise indicated, all material for this paper was collected and reared by me.—

be G..C,

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 7

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

do not occur throughout the plants’ ranges, the distribution of the
moths being affected by other factors. In all, 34 species of umbels
were examined. For the most part these food plants were situated
along highways and so were readily accessible. I have used this
method of roadside collecting for many years and have found that
larvae can thereby be collected rapidly and with much saving of time.

On the westward journey I made observations and collections at
several localities, but it was not until we reached Washington and
Oregon that serious collecting really began. A brief discussion of
some of the localities, with illustrations of the habitats and some of
the food plants, seems appropriate in order to acquaint the reader
with the diverse conditions under which these umbelliferous plants
flourish.

The Umbelliferae are found throughout the continent from the
seashore to the high altitudes of the Rocky, Cascade, and other moun-
tains, in swamps, and in the deserts and prairies. By following the
advent of spring from the lowlands to the high altitudes, it is possible
to enjoy profitable collecting from March to August. Even in the
lowlands of the Midwest and eastern United States and Canada,
certain species of larvae will be found well into August.

After crossing the Rocky Mountains, going west, one encounters
the Intermountain Area between the Rocky and Cascade Mountains.
Throughout this inland empire Umbelliferae abound, and it is here
that we find a vast number of species of Lomatium. One species of
Lomatium is found as far east as Missouri; the rest are western.
Perhaps the commonest species is L. dissectwm, which covers vast
areas from Alberta to British Columbia, Montana, Idaho, Washing-
ton, and Oregon to Colorado and California. This is host to Depres-
saria leptotaeniae.

In central Washington and Oregon, desert conditions, which ex-
tend north into southern British Columbia and south into California,
prevail. In somewhat restricted, sandy habitats within this area will
be found Pteryxia terebinthina foeniculacea, the host to D. yakimae
(pl. 1, figs. 1, 2). In Ten Sleep Canyon, Wyo., another variety of
this plant, P. t. calcarea, is host to Depressaria pteryxiphaga, described
on page 16.

Continuing west to the eastern slopes of the Cascade Mountains
one encounters a great array and succession of umbelliferous plants
that are hosts to a considerable number of species of Depressaria
and Agonopterix. One of the commonest species of these plants is
Cicuta occidentalis, which is found in marshy areas and along creeks,
rivers, and irrigation ditches. On plate 2, figure 1, is illustrated a

NO. 7 MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 3

typical habitat at Rock Creek, Oreg., where Oregon State Highway
82 crosses it, in which the Cicuta grows in association with grasses,
tule, and Typha. This particular stand of C. occidentalis was heav-
ily infested with D. julella. On plate 2, figure 2, are illustrated unin-
fested specimens of this plant at Carlton, Wash.

As one leaves the desert and arid spaces of the Intermountain
Area and ascends the eastern slopes of the Cascade Mountains he
passes through the Timbered Transition Zone. In this zone, usually
on the open, rocky slopes, Lomatium triternatum macrocarpum, host
to D. betina, is found. Also in this zone one encounters Osmorhiza
occidentalis and O. chilensis. Both of these species are attacked by
Agonopterix rosaciliella, but I have never found a Depressaria at-
tached to either of these plants. O. occidentalis is found most abun-
dantly at altitudes of 5,000 to 6,000 feet, and O. chilensis usually
grows at lower altitudes and in the Humid Transition Zone of the
west side of the Cascades. O. chilensis is not normally infested ; in
fact, the only infestation I have ever found was at Billy Goat, Okano-
gan County, Wash., where the host was attacked by Agonopterix
rosaciliella.

The alpine meadows, which one encounters after passing through
the foothills of the Cascade Range and the high ridges above them,
provide the habitats for a great variety of umbels. At Harts Pass,
which forms the boundary between Whatcom and Okanogan Coun-
ties, Wash., and Slate Peak, in the former county (pl. 3), there are
no fewer than twelve species of Umbelliferae. These are: Osmo-
rhiza occidentalis, O. chilensis, Heracleum lanatum, Angelica arguta,
Ligusticum purpureum, Lomatiwm dissectum, L. brandegei, L. geyeri,
L. ambiguum, Angelica lyalli Wats., Lomatium gormani (Howell)
C.& R., and L. triternatum (Pursh) C. &. R., the last three being
recorded by Muenscher.? In addition, L. angustatum, from which I
have reared D. angustati, will be found on some of the high ridges
such as Skyline Ridge, Mount Baker District, Whatcom County,
Wash.

Proceeding westward, after passing through the alpine meadows
and the Arctic Highland ridges, we enter the forested Hudsonian,
Canadian, and Humid Transition Zones in that order. In the first
two there is little of interest for one devoted to the study of the
insects being discussed, but in the Humid Transition there is a wide
variety of Umbelliferae.

Oenanthe sarmentosa (pl. 5, fig. 1), the host of D. nervosa, is

2 Muenscher, W. C., The flora of Whatcom County, State of Washington,
Pp. 108-110, 1941.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

abundant in practically every swale, swamp, marsh, and roadside
ditch, and although nervosa is not found throughout the range of
the host it is widespread in Oregon and Washington and undoubtedly
will be found in British Columbia. I obtained nervosa in quantity
on the American side of the international boundary at Blaine, Wash.
On the dry prairies of the Humid Transition Zone, on the dry,
gravelly slopes of some of the San Juan Islands, and along the sea-
shore at Bellingham and Birch Bays, Lomatiwm utriculatum, host
to D. besma, and L. nudicaule abound. Sium suave, Cicuta douglasii,
and Osmorhiza chilensis are also found abundantly, C. douglasii
being an occasional host of D. nervosa. In this zone also Angelica
arguta, A. lucida, and A. hendersonti are encountered. A. lucida
grows in the marshy, tidewater areas of the Siletz River, Oreg., and
A. henderson (pl. 4, figs. 1, 2) is found along the rocky bluffs of
the seashore of the Oregon coast. Both of these plants, together
with Ligusticum apufolium and Contoselinum chinense, are hosts to
A, rosaciliella and A. oregonensis.

Throughout all these zones, with the exception of the Arctic, we
are apt to find Daucus carota, Heracleum lanatum, and Conium
maculatum. H. lanatum is frequently infested with D. heracliana,
and the other two sometimes show injury, but I have not yet reared
an oecophorid from either one.

The species of Depressaria adhere rather closely to a pattern of
attack in which the young umbels are webbed by the larva or sev-
eral larvae. Pupation takes place in a hollow stalk of the host or in
debris at or near the base of the plant. Some exceptional variations
are discussed under the pertinent species.

On plate 5, figure 1, are figured characteristic examples of umbels
of Oenanthe sarmentosa in which the rays are drawn together and
the inflorescence is distorted by the larvae of Depressaria nervosa.
In figure 2 of the same plate is an illustration of an umbel of Loma-
tium dissectum, multifidum webbed and damaged by larvae of D.
leptotaeniae. When this photograph was taken the fruits were matur-
ing, but the typical damage is well illustrated.

The larvae of Agonoptcrix species are chiefly leafrollers, but occa-
sionally they web the rays and feed in the fruits. As far as I know
the larvae never pupate in a hollow stalk.

Above I have discussed the more important plants, of a rather
limited region of the continent, attacked by larvae of the two genera
under consideration and have indicated some of the places where
they may be found. Obviously I have not shown the extent of the
entire ranges of the plants, nor have I recorded all the exact localities
from which the plants have been collected.

NO. 7 MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 5

LIST OF LOCALITIES WHERE FOOD PLANTS (UMBELLIFERAE)
OF DEPRESSARIA AND AGONOPTERIX WERE COLLECTED

[“x” indicates infestations of larvae and/or collections that produced adults;

”

“o” indicates no occurrence of the insects. ]
Localities Food plants * Larvae Adults
1. Oelwein, Iowa Zizia aurea (L.) Koch 3 to)
2. Quinn, S. Dak. Lomatium sp. x )
3. Ten Sleep, Wyo. Pteryxia terebinthina calcarea
(M. E. Jones) x Bs
4. Deer Lodge, Mont. Lomatium dissectum multifidum
(Nutt.) M. & C. x x
5. Alberton, Mont. Lomatium ambiguum (Nutt.)
CA aR: x x
Lomatium macrocarpum (H. &
ALY) Cibo x me
6. Hooper, Wash. Cicuta occidentalis Greene oO 0
Comum maculatum L. 0 0
7. Washtucna, Wash. do. fe) oO
8. 8 miles west of Moses Pteryxia terebinthina foenicu-
Lake, Wash. lacea (T. & G.) Math. x x
9. to miles east of Burke, do. 5 x
Wash.
10. Pomona, Yakima Lomatium sp. fe) fe)
County, Wash.
11. Cliffdell, Kittitas Lomatium. triternatum macro-
County, Wash. carpum (C. & R.) Math. 52 x
Osmorhiza occidentalis (Nutt.)
Torr. Oo 0)
12. Sawmill Flat, Kittitas Lomatium triternatum macro-
County, Wash. carpum x x
13. Marietta, Whatcom Oenanthe sarmentosa Presl. @ 0
County, Wash.
14. Lake Samish, Whatcom do. a ae
County, Wash.
15. Bellingham, Wash. do. (0) 0)
(Numerous observations )
16. Lawrence, Whatcom Oenanthe sarmentosa x x
County, Wash.
17. Blaine, Wash. do. x Xx
Heracleum lanatum Michx. 0 )
18. Ferndale, Wash. Oenanthe sarmentosa (a) oO
19. Birch Bay, Whatcom do. 58 Be
County, Wash. Lomatium nudicaule (Pursh)
Cgc: O 9)
20. Hamilton, Skagit Oenanthe sarmentosa x x
County, Wash.
21. Vogler Lake, Skagit do. O 0
County, Wash. Osmorhiza chilensis Hook. &
Arn, 0 0
22. Toad Lake, Whatcom do. 0 oO
County, Wash. Heracleum lanatum 0 0
Oenanthe sarmentosa x x

* Botanical authority is given only after the first occurrence of the name.

SMITHSONIAN

Localities

Fazon Lake, Whatcom

County, Wash.
Olympia, Wash.
Centralia, Wash.
Chehalis, Wash.
Kelso, Wash.
Oregon City, Oreg.

Molalla, Oreg.

. Wilhoits Springs, Oreg.

Clackamas River, Oreg.

Depoe Bay, Oreg.

Hogarty Creek, Oreg.

Whale Cove, Depoe
Bay, Oreg.

Beverly Beach, Oreg.

Agate Beach, Oreg.

Siletz River, Oreg.

. Toledo, Wash.

Tumwater, Wash.

. 3 miles west of Monroe,

Wash.
Tye River, U. S. High-
way 2, Wash.
Rayrock Springs, Ste-
phens Pass, Wash.

. Winton, Wash.

Azwell, Wash.

Carlton, Wash.

Billy Goat, Okanogan
County, Wash.

. 2 miles south of Billy

Goat, Wash.

Eight Mile Creek Guard
Station, Okanogan
County, Wash.

Sherman Guard Station,
Okanogan County,
Wash.

Winthrop, Wash.

. Robinson Creek, Oka-

nogan County, High-
way 16, Wash.

MISCELLANEOUS COLLECTIONS

Food plants *
Oenanthe sarmentosa

Cicuta douglasti (DC.) C. & R.

Oenanthe sarmentosa

do.

do.
Daucus carota L.
Oenanthe sarmentosa
Osmorhiza chilensis
Heracleum lanatum
Oenanthe sarmentosa

do.
Ligusticum apiifoliwn

A. Gray

( Nutt.)

Angelica hendersonti C. & R.

Conioselinum chinense
ISIE
Daucus pusillus Michx.
Oenanthe sarmentosa
Heracleum lanatum
Oenanthe sarmentosa
Conioselinum chinense
Oenanthe sarmentosa
do.
do.
Angelica lucida (L.)
Oenanthe sarmentosa

do.
Angelica arguta Nutt.

do.
Heracleum lanatum
Angelica arguta
Osmorhiza chilensis

Angelica canbyi C. & R.

Cicuta douglasii
Heracleum lanatum
No umbels found
Cicuta occidentalis
Osmorhiza occidentalis

Osmorhiza chilensis

Angelica arguta

do.

Cicuta occidentalis
do.

(L.)

Larvae

moOoodOCOCOoOOXxXx Oo

ital

) ) (2) bs} te fe) ey fe) ) ) fe}

tat (ey CSS) To) ey tle) ()

* Botanical authority is given only after the first occurrence of the name.

VOL. I17

Adults

rat (©) () (©) Ie) (©) te} (6) ) pal

=) ©) Te) ht ye) we) EN ) fe) ) fe)

tak tebe) (oe) (e) (el Ey Te} fe)

m”

NO. 7 MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 7

Localities Food plants * Larvae Adults
52. Harts Pass, Okanogan Lomatium brandegei (C. & R.)
County, Wash. F, Macbr. oO
Lomatium geyeri (S. Wats.)
(Stoo IRR. Oo (6)
Lomatium ambiguum re) fe)
Osmorhiza occidentalis oO fa)
Angelica arguta re) fo)
53. Lower Harts Pass, Oka- Heracleum lanatum , oO 0
nogan County, Wash. Angelica arguta ra) ra)
Ligusticum sp. fe) fe)
54. Slate Peak, Whatcom Lomatium ambiguum x x
County, Wash. Lomatium geyeri re) oO
Lomatium brandegei Br 3%
Lomatium dissectum (Nutt.)
Math. & C. re) ra)
55. Ingalls Creek, Blewett Angelica arguta rs) ra)
Pass, Wash.
56. Peshastin Creek, Blew- do. ro) oO
ett Pass, Wash.
57. Touchet, Wash. Daucus carota fo) 0
58. Freewater, Oreg. Cicuta occidentalis oO 0
59. Langdon Lake, Oreg. Angelica arguta fo) oO
60. 16.5 miles north of Elgin, do. fo) 0
Oreg.
61. 10 miles north of Elgin, do. ra) 0
Oreg.
62. Minam, Oreg., Wallowa Cicuta occidentalis oO 0
River
63. Rock Creek, Highway do. x x
82, Oreg.
64. Joseph, Oreg. Heracleum lanatum 0 ty)
65. Wallowa Lake, Oreg. do. Oo 0
Angelica arguta fo) 0)
66. Aneroid Lake Trail, do. ) fy)
Oreg. Osmorhiza occidentalis fo) oO
Ligusticum sp. fo) fa)
Heracleum lanatum fo) fo)
67. Oregon City, Oreg. Oenanthe sarmentosa oO oO
Osmorhiza chilensis C0) (0)
68. Bridal Veil Falls, Oreg. Lomatium angustatum (Coult.
& Rose) St. John x 0
Lomatium triternatum macro-
carpum (a) O
Lomatium dissectum multifidum 0 0
Heracleum lanatum x x
Angelica sp. fe) Ce)
Daucus carota (injury only de-
tected) oO oO
Oenanthe sarmentosa fe) oO
Osmorhiza chilensis (0) (0)
69. Oneonta Gorge, Oreg. Lomatium angustatum (0) oO
70. Lake Padden, Whatcom Cicuta douglasii x x
County, Wash. Sium suave Walt. Cy) ty)

* Botanical authority is given only after the first occurrence of the name.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Localities Food plants * Larvae Adults
71. Waterville, Wash. Cicuta occidentalis x x
72. Coeur d’Alene Lake, Lomatiwm dissectum multifidum o o
Idaho
73. Denna Mora Creek, Ligusticwm canbyi C. & R. oO 0
U. S. Highway 10, Angelica arguta ra) oO
Mont.
74. Clinton, Mont. Sium. suave oO oO
75. McDonald Pass, Mont. Angelica arguta x x
Heracleum lanatum
Osmorhiza occidentalis (0) fo)
76. Jackson Lake, Wyo. Ligusticum filicinum S. Wats. (0) 9)
77. Togwotte Pass, Wyo. do. Be (0)
78. 25 miles southeast of Lomatiwm sp. () 0
Lander, Wyo.
79. 13 miles east of North Cicuta maculata L. x x
Platte, Nebr.

80. Lucas, Iowa do. x x

* Botanical authority is given only after the first occurrence of the name.

As pointed out at the beginning of this discussion, the object of
the field work undertaken was to endeavor to induce larvae to accept
substitute foods, and to rear the moths. On page g is a chart in which
I have tabulated the food plants and the species of Depressaria and
Agonopterix that have been reared to date. It will be noticed at once
that the differences in feeding habits between the species of the two
genera follow different patterns. The three species of Agonopterix
are definitely polyphagous, one species feeding on no less than 14
species of umbels. The species of Depressaria, on the other hand,
show a distinct monophagous tendency, although three species feed
on more than one plant. Even so, the range of acceptance is very
narrow and, indeed, though not lethal, may be accidental.

The first attempt to induce larvae to feed on substitute food was
made in 1933 when larvae of Depressaria multifidae were offered,
in place of Lomatium grayt, the normal food, L. dissectum multifidum,
which they refused. In 1946,? however, a series of moths was reared
from L. columbianum which are indistinguishable from a long series
of multifidae.

In the summer of 1950 numerous attempts at substitution of foods
were made, but without success. Larvae of D. betina found on L.
triternatum macrocarpum, the normal food plant, were offered Os-
morhiza chilensis. Most larvae refused this substitute, but some ate
it voraciously ; all died. Although L. triternatum macrocarpum is
the normal food, occasionally scattered larvae from which moths
were reared were found on L. dissectum and L. columbianum. All

8 Clarke, J. F. Gates, Journ. Washington Acad. Sci., vol. 37. pig w1o4zs

NOT 7, MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 9

these species of Lomatium are frequently found growing in close
association.

Depressaria nervosa was found in great abundance in the extreme
western parts of Whatcom and Skagit Counties, Wash., and less

cinereocostella

whitmant

schellbacht

heracliana
juliella
leptotaeniae
yakimae
multifidae
angustati
constancet
betina
moya
besma

eleanorae
togata

DEPRESSARIA
angelicivora
pteryxiphaga
nervosa

AGONOPTERIX
oregonensis
rosaciliella
muricolorella

thustra
armata

Angelica arguta
hendersonii
lucida x
Cicuta douglasii x
maculata x
occidentalis Sto oe
Contoselinum chinense x
Daucus carota ?
Eryngium vaseyt x
Heracleum lanatum Se
Ligusticum apiifolium xk
Lomatium ambiguum xs
angustatum x
brandegei x
californicum x
carutfolium x
columbianum x x
dissectum x
dissectum multifidum x
grayi x x
macdougali be
macrocarpum x x
marginatum x
nudicaule x
triternatum macrocarpum x x
utriculatum x Be
vaginatum x
Oenanthe sarmentosa Sie) acre
Osmorhiza chilensis
occidentalis x
Pastinacea sativa x
Pteryxia terebinthina calcarea 52
terebinthina foeniculacea x
Sanicula bipinnata
bipinnatifida
laciniata
nevadensis
tuberosa

a
ra
“mA

“

ARK KK

commonly in one locality in Oregon, feeding on Oenanthe sarmentosa.
Larvae of this moth feed on a species of Oenanthe in Europe and,
undoubtedly, in Asia. At Fazon Lake and Lake Padden, both in
Whatcom County, Wash., I found a few larvae on Cicuta douglasii,
which was quite acceptable to them, and they completed their meta-
morphosis. The food plant at Lake Padden was growing intermingled
with Sium suave, which the larvae did not infest and which was
refused by them when offered as a substitute. Larvae of nervosa were

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

offered umbels of Angelica arguta and Cicuta occidentalis, which also
were refused.

At Slate Peak, Whatcom County, Wash., a large quantity of larvae
of an undescribed species of Depressaria was collected on Lomatium
brandegei. Six larvae were offered Oenanthe sarmentosa, six more
Osmorhigza chilensis, but all refused these substitutes and died. One
larva accepted L. angustatum, from the Columbia River Gorge, Oreg.,
but died.

Other larvae collected at Slate Peak, feeding on L. geyeri, were
offered L. angustatum and O. chilensis, which were refused, and all
the larvae died.

Larvae of D. juliella were found on the usual food plant, Cicuta
occidentalis, where Oregon State Highway 82 crosses Rock Creek,
Oreg., a tributary of the Wallowa River. Four of these larvae were
offered Angelica arguta, but they refused to eat it and remained
inactive in the rearing tin.

Only one substitution in the genus Agonopterix was attempted and
that for A. rosaciliella. Larvae collected on Osmorhiza chilensis were
given O. occidentalis, which they readily accepted. All produced
moths.

We can therefore infer that in Depressaria a high degree of host
specificity exists and that, although larvae can develop on more than
one food plant when once established, no tolerance exists for sub-
stitutes in diet. It is equally clear that no host specificity exists in
Agonopterix and that there is a correspondingly high degree of toler-
ance for substitutes.

AGONOPTERIX MURICOLORELLA (Busck)
Depressaria muricolorella Buscx, Proc. U. S. Nat. Mus., vol. 24, p. 741, 1902.

Type—U.S.N.M. No. 6125.

Type locality.—Golden, Colo.

Food plants —Lomatium grayi C. & R.* and L. macrocarpum.

Remarks.—Of this species I now have two specimens from Alber-
ton, Mont., reared from Lomatium macrocarpum, recorded here for
the first time. Both specimens are males and emerged July 11-13, 1950.
The larvae were found feeding in the immature fruits of the host.

4 Authorities for botanical names are given only for those names not appear-
ing in the list, p. 5 e¢ seq.

SMITHSONIAN MISCELLANEOUS COLLECTIONS WAONLs Baba WOE ZA Tele 2b

Fig. 1, upper: Pteryxia terebinthina foeniculacea growing in association with
grasses, Artemisia, and Chrysothamnus 8 miles west of Moses Lake, Wash.
Fig. 2, lower: Pteryxia terebinthina foeniculacea, same locality.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 7, PL. 2

Fig. 1, upper: Cuicuta occidentalis growing in association with tule, grasses, and
2. lc

Typha at Rock Creek, Oreg., at the junction with State Highway 82. Fig

o, 2, lower:
Cicuta occidentalis, uninfested, at Carlton, Wash.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS ait INOS 7, RES 3

een

am “ aes (ia

Fig. 1, upper: Slate Peak, Whatcom County, Wash. Four species of Lomatiwm
are found on the slopes shown in the center of the photograph. These are L. ambi-
guum, L. brandegei, L. dissectum, and L. geyeri. Fig. 2, lower: The same locality

as above but farther east. On the steep shale slopes in the upper part of the illustra-
tion the same four species of Lomatium listed above are found.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 7, PL. 4

Mig. 1, upper: Angelica hendersonti growing along the rocks of the Oregon coast
at Finistere Lodges, Depoe Bay. On the ground above Conioselinum chinense and
Heracleum lanatum are abundant. Fig. 2, lower: Angelica hendersonii in the same
locality as above.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO eizin INO 7; aPlee

Fig. 1, upper: Ocnanthe sarmentosa growing in association with ferns, Lysichi-
ton, Mimulus, and grasses at Lawrence, Whatcom County, Wash. Note three dis-
torted umbels in center of photograph which are infested. Fig. 2, lower: Lomatium
dissectum multifidum from Pullman, Wash. Note the characteristic damage and
webbing done by larvae of Depressaria leptotaeniae.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO, Liz, NOs wR IEG

Figs. 1-1b, Depressaria angelicivora, new species: 1, Right harpe; Ia, aedeagus;
1b, anellus. Figs. 2, 2a, Depressaria angelicivora, new species: 2, Female genitalia
with ovipositor and bursa copulatrix removed; 2a, signum. Figs. 3-3b, Depressaria
pteryxiphaga, new species: 3, Right harpe; 3a, aedeagus ; 3b, anellus. Figs. 4, 4a,
Depressaria pteryxiphaga, new species: 4, Female genitalia with ovipositor and
bursa copulatrix removed; 4a, signum. Figs. 5-5b, Depressaria armata, new species :
5, Right harpe; 5a, aedeagus; 5b, anellus.

NOs 7, MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE iit

AGONOPTERIX OREGONENSIS Clarke
Agonopterix oregonensis CLARKE, Proc. U. S. Nat. Mus., vol. 90, p. 65, I94I.

Type.—U.S.N.M. No. 52079.

Type locahity.—Salem, Oreg.

Food plants—Angelica hendersoni, A. lucida, Eryngium vaseyi
C.& R., Ligusticum apitfolium, Lomatium caruifolium (H. & A.)
C. & R., L. marginatum (Benth.) C. & R., L. nudicaule (Pursh)
C.& R., L. utriculatum (Nutt.) C. & R., Oenanthe sarmentosa,
Sanicula bipinnata H. & A., S. bipinnatifida Dougl., S. laciniata H. &
A., S. nevadensis S. Wats., and S. tuberosa Torr.

Remarks.—Only four of the food plants listed above, A. hender-
soni, A. lucida, Ligusticum apiifolium, and Oenanthe sarmentosa,
are recorded here for the first time; the others, recorded previously
by me,° are entered for completeness.

The localities for the specimens reared in 1950 are as follows:
Oregon: Depoe Bay, 5 33, 2 99 (July 26-28, 1950) ; Siletz River,
2 miles east U. S. Highway rot, 2 J/g (August 3, 1950). Washington:
Lawrence, Whatcom County, 2 (July 22, 1950) ; Toad Lake, What-
com County, J (July 21, 1950).

AGONOPTERIX ROSACILIELLA (Busck)
Depressaria rosaciliella Buscx, Proc. U. S. Nat. Mus., vol. 27, p. 763, 1904.

Type.—U.S.N.M. No. 7815.

Type locality—‘‘Camp Watson,” Oreg.

Food plants—Angelica arguta, A. hendersonii, Conioselium chi-
nense, Ligusticum apufolium, Oenanthe sarmentosa, Osmorhiza
chilensis, and O. occidentalis.

Remarks.—In my revision of this family (p. 84) ® I recorded the
food plant of this species as Osmorhiza occidentalis and stated that
the larvae of rosaciliella were not found on O. chilensis. On a preced-
ing page of the present paper I have indicated that the species is
found on both plants. I have before me 19 specimens of rosaciliella
reared during the summer of 1950 from the above list of hosts. These
show a wide degree of variation, including specimens that match the
Blue Mountains material recorded previously (p. 84) ® and also moths
that are of the exact color and contrasting markings as the series

5 Clarke J. F. Gates, Journ. Washington Acad. Sci., vol. 37, p. 3, 1047.

6 Clarke, J. F. Gates, Revision of the North American moths of the family
Oecophoridae, with descriptions of new genera and species. Proc. U. S. Nat.
Mus., vol. 90, pp. 33-286, pls. 1-48, 1941.

I2 SMITHSONIAN MISCELLANEOUS. COLLECTIONS VOL. 117

of 90 specimens I recorded from Skyline Ridge, Whatcom County,
as A. r. echinopanicis, which were reared on Echinopanax horridum
(p. 86) and which I described as a food-plant race of rosaciliella.
There is no doubt that all these color variations represent entities of
a single species. There appears to be no justification for the racial
designation of echinopanicis, although those who wish to use this
name for the population feeding on E. horridum may do so for this
contrastingly marked form.

The localities for the moths reared in 1950 are as follows: Oregon:
Clackamas River, 15 miles east of Estacada, 2 df (July 28, 1950),
2 (July 27, 1950); Depoe Bay, 3 dg (July 27-August 2, 1950),
5 92 (July 28-August 2, 1950). Washington: Billy Goat, Okanogan
County, 2 dd, 4 92 (July 29-August 2, 1950) ; Lawrence, Whatcom
County, 2 (July 24, 1950); Rayrock Springs, Stephens Pass, 9
(August 2, 1950).

DEPRESSARIA CINEREOCOSTELLA Clemens

Depressaria cinereocostella CLEMENS, Proc. Ent. Soc. Philadelphia, vol. 2,
p. 422, 1864.

Type.—In the Academy of Natural Sciences of Philadelphia.

Type locality —“Virginia.”

Remarks.—In my revision I listed three food plants for this species,
Carum carvi L., Sium lineare Michx., and Ligusticum scoticum L,
It is impossible to check the identities of the plants from which the
specimens are supposed to have been reared, but it seems certain
that there has been some confusion regarding some of the identifica-
tions of the plants made many years ago, before the Umbelliferae
were properly revised by Mathias and Constance. One of the plants
listed above, S. lineare, is a synonym of S. suave Walt., as is also
S. cicutaefolium Benth. & Hook. I have before me a series of cine-
reocostella labeled “Rf. Sium cicutaefolium,’ which I collected at
the junction of South River and U. S. Highway 50, Maryland. These
bear emergence dates of August 11-20, 1939. The identity of this
plant has not been verified by competent authority.

In August 1950 larvae of cinereocostella were encountered com-
monly in several places in Nebraska and Iowa, feeding in the umbels
of Cicuta maculata. I have before me a series of 12 males and 12
females from 13 miles east of North Platte, Nebr., and 7 males and
6 females from Lucas, Iowa. Emergence dates range from August
8-21, 1950.

The larvae of the South River specimens pupated in the hollow

NO. 7 MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE iS

stalks of the host, a habit found commonly among Depressariae. The
larvae of the Nebraska and Iowa specimens, however, pupate in a
tightly webbed umbel. Actually the rays of the umbels are left free,
but the flowers are webbed into a compact mass.

Despite these differences in habits I can find no structural or col-
orational differences that suggest specific separation.

DEPRESSARIA NERVOSA Haworth
Depressaria nervosa HawortH, Lepidoptera Britannica, vol. 3, p. 560, 1811.

Type—In the British Museum (Natural History).

Type locality—London, England.

Food plants——Oenanthe crocata L., O. sarmentosa, and Cicuta
douglasii.

Remarks.—Walsingham* recorded this species from “Southern
Oregon” in 1881, but there has always been some doubt about the
accuracy of his identification, and the name of the European nervosa
has been dropped from the North American lists. Busck ® believed
that Walsingham’s specimens were probably referable to D. juliella
Busck, and the name nervosa was omitted from Busck’s revision of
the family.*° The similarity between nervosa and juliella is apparent,
but the latter is much brighter and lighter than the former and their
food plants are different.

In England D. nervosa larvae feed in the umbels of Oenanthe
crocata and in western North America in the umbels and on the
leaves of O. sarmentosa and Cicuta douglasu. Despite these differ-
ences in the food plants there is no doubt that the American speci-
mens are referable to nervosa, although the Washington and Oregon
series might warrant subspecific designation.

I have before me a reared series of 173 specimens as follows:
Oregon: Clackamas County, Wilhoits Springs, 3 J, 3 92. Emer-
gence dates, July 25-29, 1950. Washington: Skagit County, Hamilton,
o, 4 99; Whatcom County, Birch Bay, 3 J, 6 92; Blaine, ¢; Fazon
Lake, gf, 2 99; Lake Padden, 2 J’, 9; Lawrence, 72 do, 57 99;
Samish Lake, 5 dd’, 8 99; Toad Lake, 2 ¢¢, 9. In addition to the
above there are 24 specimens deposited in the British Museum (Nat-
ural History).

The emergence dates range from July 17 to August 18, 1950. The
August dates are for specimens collected at Lake Padden and reared
from Cicuta douglas.

7 Proc. Zool. Soc. London, 1881, p. 317.
8 Proc. Ent. Soc. Washington, vol. 9, p. 91, 1908.
2 Proc. U. S. Nat. Mus., vol. 35, pp. 187-207, 1908.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

I am indebted to John Bradley, of the British Museum, who has
kindly compared some of the American specimens with the European
series in the British Museum collections and who has confirmed my
identification. The European specimens in the U. S. National Mu-
seum are lighter and more olivaceous than the moths from Oregon
and Washington, but the latter vary from olivaceous to rather bright
red although the majority are predominantly brownish red.

DEPRESSARIA WHITMANI Clarke

Depressaria whitmani CLARKE, Proc. U. S. Nat. Mus., vol. 90, p. 182, pl. 36,
figs. 200, 200a; pl. 48, fig. 286, 1941.

Type.—uU.S.N.M. No. 52083.

Type locality——Snake River, Whitman County, Wash., opposite
Clarkston.

Food plant—Lomatium macrocarpum.

Remarks.—In addition to the type series, I now have a female,
reared from the normal food plant, from Montana. The larva was
collected 3 miles east of Alberton, on U. S. Highway Io, feeding in
the fruits of the host on June 15 and the moth emerged July 7, 1950.
The food plant was growing on rocky ground in association with
pine, grasses, and balsam root. The occurrence of this species in
Montana extends the range over 200 miles east of the type locality.

DEPRESSARIA YAKIMAE Clarke
Depressaria yakimae CLarkeE, Proc. U. S. Nat. Mus., vol. 90, p. 185, 1941.

Type.—U.S.N.M. No. 52073.

Type locality—Yakima, Yakima County, Wash.

Food plant.—Pteryxia terebinthina foeniculacea.

Remarks.—When I described this species I predicted that the larva
would be found on some species of Lomatium, but I have now found
yakimae attached to Ptery-xia, a closely related genus.

I now have six reared specimens that were obtained from pupae,
the larval stage having passed by mid-June. The pupae were col-
lected at a point 8 miles west of Moses Lake, Wash., on U. S. High-
way 10, and also 10 miles east of Burke, Grant County, Wash., on
U.S. Highway to. The dates were June 17 and 21, 1950, respectively.
Moths began to emerge June 18 and the last appeared June 23, 1950.

The larva of this species spins a tough though rather loose cocoon
between the rays of the flowers where pupation takes place. This
is a departure from the usual habit for pupation in this group and

NO. 7 MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 15

is parallel only to cinereocostella among the known species. The pupa
is always found with the ventral surface down and the caudal end
elevated, sometimes almost to the vertical position.

DEPRESSARIA ANGELICIVORA, new species
Plate 6, figures 1-Ib, 2-2a

Description—Alar expanse, 25-27 mm.

Labial palpus with second segment creamy white strongly tinged
with pink inwardly, with scattered fuscous scales outwardly and the
brush suffused with fuscous; third segment fuscous sparsely irrorate
with pink scales and with pink tip. Antenna fuscous except under-
side of scape, which is pink, and underside of about one-third of
shaft, which is pink to cream color. Head and thorax whitish ocherous
with admixture of fuscous on vertex and anteriorly on thorax and
tegula. Forewing with basal third blackish fuscous shading to paler
fuscous at apex, streaked with blackish fuscous along veins in some
specimens and costa edged with pink, the whole with a washed, faded
appearance ; from costa at middle an inwardly oblique whitish ocher-
ous line to middle of cell continuing as an outwardly oblique though
more indefinite line of the same color to middorsum; from slightly
beyond inception of the above line a similar, outwardly oblique line
to apical fourth at center of wing, thence inwardly oblique to slightly
before tornus; these two lines are shaded and streaked with pale-
brownish scaling and outline a more or less diamond-shaped area
in center of wing; at end of cell a whitish ocherous spot; from
apical third of costa around termen to middorsum a somewhat broken
blackish fuscous line; cilia fuscous basally, lighter distally with some
pinkish suffusion. Hindwing whitish basally shading to fuscous
around margins; cilia pale yellowish fuscous with fuscous basal line
and suffused with pink. Legs whitish ocherous banded and suffused
with fuscous and pink. Abdomen whitish ocherous suffused with
fuscous above and with blackish fuscous longitudinal lateral band
beneath.

Male genitalia—Similar to other members of this group, but the
basal process of sacculus about as long as half the width of harpe
at base and the clasper straight, digitate.

Female genitalia—Genital plate narrow and anterior pockets shal-
low ; sclerotized section of ductus bursae very short, about one-fourth
total length.

Type.—U.S.N.M. No. 61133.

Type locality—McDonald Pass, 14 miles west of Helena, Mont.
(6,100 feet).

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Food plant—Angelica arguta.

Remarks.—Described from the type 3, 5 d' and 3 2 paratypes, all
from the type locality. Emergence dates August 9-12, 1950. Para-
types in the U. S. National Museum and the British Museum (Nat-
ural History).

The food plant of this species is widespread, but the only examples
found infested were the specimens from which this series of moths
was reared. The plants were growing in moist ground at a spring
with its resulting drainage, and were growing in association with
Heracleum, Mimulus, Epilobium, Alnus, and other species comprising
a lush growth.

The larva of this species attacks the young umbels before they
have opened completely and causes great distortion and discoloration
of the inflorescence. Pupation occurs in the leaf bracts near the
main stalk of the host. This is a departure from the usual habit of
members of this group, which almost always pupate in debris near the
base of the plant or in the hollow stalks. As many as five pupae
were found in a single bract, each larva having constructed a heavy
web before pupation.

In the key to species in my revision, this species runs to betulella
by reason of the longitudinal stripes on the ventral side of the abdo-
men. However, angelicivora belongs in the angustati-multifidae com-
plex and can be separated from all the species of this group by the
washed-out, faded appearance of the forewing. The male genitalia
of angelicivora place it nearest schellbachi, but it differs from the
latter by the shorter basal process of the sacculus, which is a little
more than half the length of that in schellbachi, and by the straight
clasper as compared to the curved clasper of schellbacht. The aedeagus
of angelicivora is more like those of leptotaeniae and yakimae. The
female genitalia are similar to thustra, but the sclerotized section of
the ductus bursae is slightly more than half as long.

DEPRESSARIA PTERYXIPHAGA, new species
Plate 6, figures 3-3b, 4-4a

Description.—Alar expanse, 18-20 mm.

Labial palpus with second segment ocherous-white, suffused with
pinkish in most specimens, marked with fuscous and pink-tipped
fuscous scales exteriorly and in the brush; third segment blackish
fuscous, apex ocherous-white. Antenna scape blackish fuscous, shaft
grayish fuscous with paler annulations. Head and thorax ocherous-
white to ocherous with a strong pinkish suffusion in most specimens ;

NOS 7, MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 17

thorax anteriorly, and tegula basally, edged with fuscous. Forewing
blackish fuscous basally shading to fuscous apically; extreme edge
of costa ocherous-white to ocherous suffused with pinkish; slightly
before middle of costa and at apical third of costa large blotches of
the same color with similar, smaller spot at base of costa; at end of
cell an ocherous-white to ocherous discal spot followed by a slender
black dash; at basal third, on vein 11, a black spot followed by an
oblique dash of the same color on vein 10; from apical third of costa,
around termen to tornus, an indistinct series of small black spots;
cilia grayish fuscous, darker basally. Hindwing whitish basally shad-
ing to pale fuscous around margins; cilia light buff to grayish around
apex with narrow, pale-fuscous subbasal line ; sometimes cilia suffused
with pinkish. Legs ocherous-white marked and banded with fuscous
and paler areas sometimes pinkish. Abdomen ocherous-white suffused
with grayish above and strongly marked fuscous beneath, especially
laterally.

Male genitalia —Similar to the angustati-multifidae group but with
straight basal process from sacculus with few dentate processes and
with distal end of clasper dilated.

Female genitalia—Genital plate narrow and strongly sclerotized
along anterior edge ; anterior pocket narrow ; ductus bursae sclerotized
for about one-third its length.

Type.—U.S.N.M. No. 61134.

Type locality.—Ten Sleep, Wyo.

Food plant.—Pteryxia terebinthina calcarea.

Remarks.—Described from the type J, 2 J and 4 2 paratypes, all
from the type locality. Emergence dates July 5-9, 1950. Paratypes
in the U. S. National Museum and the British Museum (Natural
History).

The food plant of this species is common in the Ten Sleep Canyon,
just east of Ten Sleep, Wyo., and the actual locality in which the
larvae of this species were collected was at the foot of the canyon
alongside U. S. Highway 16 at 4,600 feet. Here the plant grows
in association with Artemisia, Balsamorhiza, and grasses on the steep
slopes of the sides of the canyon.

The larva of pteryxiphaga feeds on the leaves of the food plant
and webs them together. Larvae were collected June 13, 1950, and
pupation began June 16, 1950.

A brief color description follows: Larva pale yellowish green with
dorsal and broad subdorsal longitudinal stripes gray-green ; tubercles
pale whitish green with brown at insertion of setae. Head pale

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

yellowish brown with sparse brown mottling posteriorly and the
sutures dark brown. Thoracic shield yellowish narrowly edged with
brown laterally ; on each side, dorsolaterally, a few dark-brown spots.
Anal plate yellowish green.

In my key this species runs to the yakimae-leptotaeniae couplet
but is easily distinguished from both by the pale costal blotches.
Occasional specimens of pteryxiphaga lack the pink costal edge, in
which cases the examples would run to the angustati-multifidae
couplet. In such examples the genitalia must be used for identifica-
tion. In all cases, as a matter of fact, the genitalia should be used in
determination of species of this complex.

The genitalia of this species suggest a close affinity with multifidae
on the one hand and thustra on the other. The twisted basal process
of the sacculus of thustra immediately separates pteryxiphaga from
it, and the latter can be distinguished from multifidae by the straight
basal process of sacculus. The female of pteryxiphaga can be sepa-
rated from multifidae by the narrow genital plate and shallow anterior
pocket and from thustra by the strongly sclerotized anterior edge of
the genital plate and the shorter sclerotized section of ductus bursae.

DEPRESSARIA LEPTOTAENIAE Clarke
Depressaria leptotaeniae CLARKE, Can. Ent., vol. 65, p. 87, pl. 4, 1933.

Type.—U.S.N.M. No. 44742.

Type locality—Pullman, Wash.

Food plant.—Leptotaenia dissectum multifidum (Nutt.) M. and C.

Remarks.—Larvae of this species were collected on the host on a
hillside 11 miles northwest of Deer Lodge, Mont., on June 15.
Pupation began the following day, and the moths, one male and two
females, emerged July 8-9, 1950. The food plant was growing in
association with sagebrush, balsamroot, grasses, and other prairie
plants on open ground.

This species has not been recorded previously in Montana, and the
distribution is thus extended eastward considerably.

DEPRESSARIA THUSTRA Clarke
Depressaria thustra CLARKE, Journ. Washington Acad. Sci., vol. 37, p. 15, figs.
7-74, 14, 1947.
Type—uvU.S.N.M. No. 58009.
Type locality —Gilmer, Klickitat County, Wash.

Food plants—Lomatium triternatum macrocarpum and L. am-
biguum.

NO. 7 MOTHS, DEPRESSARIA AND AGONOPTERIX—CLARKE 19

Remarks.—Originally I described this species from the first-named
food plant from a low altitude. I have before me, however, a single
specimen from L. ambiguum, from Slate Peak, Whatcom County,
Wash., 6,500 feet, which is indistinguishable from the original series.
The twisted basal process of the sacculus is characteristic and distin-
guishes thustra from all other described species.

The Slate Peak specimen emerged August 4, 1950; larva collected
July 18, 1950.

The food plant (ambiguum) of this species was found at Harts
Pass and on the rocky slopes of Slate Peak (pl. 3), but larvae were
found only at the latter place where they were observed tying and feed-
ing in the umbels.

When the supply of the original food plant became exhausted I
substituted L. angustatum and O. chilensis, but both of these sub-
stitutes were refused and the larvae died.

DEPRESSARIA ARMATA, new species
Plate 6, figures 5-5b

Description.—Alar expanse, 17 mm.

Labial palpus with second segment light buff with moderate grayish-
fuscous scaling in brush; exteriorly and interiorly shining silvery
fuscous ; third segment grayish fuscous, buff-tipped. Antenna fuscous ;
underside of scape and proximal third of shaft shining silvery
fuscous. Head and thorax buff mixed with grayish fuscous; anterior
edge and posterior tip of thorax and basal third of tegula grayish
fuscous. Forewing grayish fuscous becoming considerably lighter
toward apex; at basal third, in cell, a longitudinal blackish dash fol-
lowed, at end of cell, by a blackish-edged whitish-buff spot ; at three-
fifths of costa an obscure blackish subquadrate spot ; from apical fifth
of costa, around termen to tornus, a series of obscure blackish spots ;
cilia grayish with a darker basal band. Hindwing very pale basally
shading to light grayish fuscous at apex; cilia grayish with darker
subbasal band. Legs whitish buff shaded and banded with grayish
fuscous.

Male genitalia—Basal process armed with fine, spinulate spines ex-
cept at base where they are shorter and coarser. Anellus very broad,
broader than long.

Female genitalia— Female unknown.

Type.—U.S.N.M. No. 61135.

Type locality.—Slate Peak, Whatcom County, Wash., 6,500 feet.

Food plant.—Lomatium. brandeget.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Remarks.—Described from the unique type male. The adult
emerged August 4, 1950.

This species is nearly related to angustati, and keys to the same
place in my key, but there are abundant points of difference between
them. The costal spot and the lack of any transverse fasciae on the
forewing of armata immediately distinguish it from angustati. The
basal process of the sacculus of armata is turned away from the cucul-
lus and that of angustati is turned toward it. The habits of the two
are also different, the larva of angustati feeding in the leaves and the
larva of armata inhabiting the umbels.

The larvae of armata were found abundantly at between 6,400 and
6,700 feet, being absent at Harts Pass at 6,200 feet and absent above
6,700 feet.

Although a considerable quantity of larvae were collected, only one
moth was obtained owing to lack of sufficient food plant and to sub-
stitution attempts. Larvae were offered Oenanthe sarmentosa and
Osmorhigza chilensis, which they refused, and all died. Others were
offered L. angustatum, but only one accepted this substitute and all
died.

sui
A!

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 8

THE SAND CRAB EMERIVTA
TALPQIDA (SAY) AND SOME
OP acre ha tIv ES

BY
R. E. SNODGRASS

(PusiicaTion 4086)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
APREE 15,1952

The Lord Galtimore Presa
BALTIMORE, MD., U. 8 As

THE SAND CRAB EMERITA TALPOIDA (SAY)
AND’ SOME, OF ITS ‘RELATIVES

By R. E. SNopGrass

CONTENTS

Pace
MER OCUICETOtecccre te ocicr a. o ciede cates eine Pouadcrare sacheve SorSusieie evens euese seiner s etek I
le General stouctune of mera ntalpotdd aasieon saeco aces eel: 3
Eh The head andiats ‘appendages 5220.2. cate fa. Se Sena Siaate eels 6
AR pte ove VGC Reet ie eee ena oe COMET be RIC CRIES Cuno nae PRL arom bce bee 7
IPO HEVESEALIES ct ac eys ah ia as cotnich = usin) cic ane eaSers buck ish Sede 9
“Whe tikst: ATCEHMNAG we /a.csrstin/o cheyerevensieueveleret ese haereretouer versie aye ereeaes wy 9
sMhessecondpantennaeictie.) = dere aetie sherincrs er naicieeneiceierieric 9
NTS NiTePera nie sls fa t0) @7b. cee BRA MORSE cicay nN ripen Mich cand Sear Sian) 13
ive Lhe enathothoracic -appendages 2. 25. is. oslo os wcjeetels a bloidel dees 17
AW sSnaarirabl ESR, 68 a4 Ag nod He Bthoa rien hadi. ce Cad od one Bo omidcn ola 17
AMIS Kiker ii bollETOe as Meare ncaa a5 oe Sets Acero nice Die eera aot hoes Ofc 18
ANite Gavanls! subdue aaah ooe pas boo deo econ oe som OcC oon UocoGT 18
PMO her ENS Ee Toa TNE INES Sta cree sas Sea tegen ernie ayay ss x Tone Oe iaey iateie senses She 18
‘TPhetsecond sawed sacenes s5s.u oss Coe Cominat tes sine scien ome 18
Aloe daigelseemalirsls So KeuobooacododcsdtooosccudouoodnOuDs at 20
mbhie! ecg) Bie. A. ela tr, wits crete eee ade Me a alee ae he a eye Pela 20
Wi hel respiratory | Systema x.a5 bint o-. dekelgeis och ls eerie yele nen eie ete ole 24
VI. The abdomen, or pleon, and its appendages.............eeeeeeeeee 27

VII. Comparison of Emerita with other members of the Hippidae and
Tyltie LHe mA blinerGaen acre ceitelcc soiree citi eeso oh hekein ert e 29
Explanation of lettering on the figures..........-+-.seeee eee eee 33
RYN Scab SA Ut Sen OR a eA nA is Hike le Bei cheap chore cic Aig Olea re arg 33

INTRODUCTION

Emerita talpoida (Say) of the Atlantic coast of the United States,
and several related species found elsewhere, are small anomuran
crabs of the family Hippidae, which are of particular interest because
of their total adaptation to a special way of living and to what is
probably, among the arthropods, a unique method of feeding. The
Emeritas bury themselves with the head end up, “facing” the ocean,
in the wet sand of sloping beaches covered by the wash of the waves.
Here they extend their long, plumose antennae to collect from the
descending water over them whatever food material may be carried
in suspension, which is then transferred to the mouth parts by a

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 8

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

ventral flexing and folding of the antennae, which in turn are en-
closed by the large platelike meropodites of the third maxillipeds.
The mandibles are much reduced and are entirely functionless as
organs of feeding. The antennal method of food gathering by Emerita
was first described by Weymouth and Richardson (1912) for the
west-coast species analoga.

Smith (1877) in his study of Emerita (Hippa) talpoida of the
Atlantic coast did not recognize the feeding function of the antennae,
which he thought were used for cleaning the legs ; finding the stomach
full of sand, he concluded that the crabs swallowed sand for what-
ever nutritive matter it might contain. The essential likeness in struc-
ture of the several Emerita species, however, can leave no doubt
that they all feed in the same manner. Their habits alone would
suggest that the antennae are food-collecting organs, and the mouth
is so closely shut in by the maxilliped plates that a direct ingestion of
sand would hardly be possible.

It is surprising to find that no other members of the hippid family
are equipped for antennal feeding, though they are all sand burrowing
in their habits. In the two other principal genera of the family, Hippa
and Mastigochetrus, the mandibles are even more reduced than in
Emerita, but the second antennae have short, simple flagella in no way
adapted to food gathering. In both these genera, however, the inner
lobes of the first maxillae are developed into a pair of strong, toothed
jaws closing beneath the mouth. Evidently, then, the functional loss
of the mandibles by the hippids has been compensated in two ways:
in Hippa and Mastigocheirus by the formation of substitute jaws
from the maxillulae; in Emerita by the development of the antennal
flagella into food-gathering organs. One is tempted to wonder why
the adult hippids lost their mandibles in the first place, and how they
fed while making adjustments to their present structure. The nature
of the food and the feeding method of Hippa and Mastigocheirus,
however, are still unknown. The members of the related family
Albuneidae, which likewise live in sand, have well-developed, func-
tional mandibles, and strongly chelate first pereiopods.

Both Emerita talpoida and E. analoga are commonly found in great
numbers between tide marks on sandy beaches, where they will be
covered by the inflowing waves. As the crabs are exposed by the
receding water they quickly back themselves into the wet sand in a
slanting position with the ventral surface toward the ocean. Here
they await the return wave, and when the downflow begins they
spread the antennae out before them against the current. According
to the tides, the crabs move up or down the beach. MacGinitie (1938)

no. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 3

says of EF. analoga that a “general migration with the tide takes place
in mass movement” and is done so quickly that to an observer “it
appears that a portion of the beach comes up, moves either up or
down, and then disappears. ... On a gently sloping beach this migra-
tion usually occurs at a time when the sand crab colony is covered
with from two to six inches of water.” When the movement of the
water begins to slacken, the crabs quickly orient themselves facing
the ocean and dig into the sand. Wharton (1942) says of E. talpoida
that the crabs “will follow a shallow wave towards the water and a
deep wave up the beach.” The usual stimulus for proper orientation,
according to MacGinitie, is the water flowing over the crabs from
behind, and if a stream of water from a hose is made to flow over
them away from the ocean, “they will immediately reverse their
position in the sand and face toward the land, even if they must move
uphill to do so.’ Under natural conditions, MacGinitie says, “the
stimulus of the current appears to be much stronger than the stimulus
of sight,” though when the current is at a minimum, if a large
black screen is placed between the crabs and the ocean, they will face
landward.

The mating of the sexes of Emerita analoga on the California coast
is said by MacGinitie (1938) to take place late in spring and early
in summer. The males, generally several at a time, bury themselves
in the sand at the side of a female and attach to her under surface
packets of spermatozoa in a thick adhesive mucus. According to
Wharton (1942), the very young males of E. talpoida, with a car-
apace length of 3 mm., which appear in August at Beaufort, N. C.,
are sexually mature, and attach themselves to year-old females for
mating, as many as seven of the miniature males having been found
on a single large female. The spermatophores are attached between
the coxopodites of the fourth legs of the female. From the time of
their first appearance in August until the following June, the small
males increase in length of the carapace from an average of about
3 mm. to about 7 mm.

I. GENERAL STRUCTURE OF EMERITA TALPOIDA

- The material on which the anatomical part of this paper is based
has been furnished by the Division of Marine Invertebrates of the
United States National Museum.

A specimen of Emerita talpoida as ordinarily seen (fig. 1 B), when
freshly dug up out of the sand, or in museum collections, has the
general appearance of a small, symmetrically egg-shaped object, the

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

largest females being about 30 mm. in length, the males half as
long. A pair of short antennules (Ant) and two slender eyestalks
project from the anterior end, and on the sides at the posterior end
are two paddlelike uropods (Urpd). The rest of the appendages are
closely folded against the underside of the body, where the legs,
except those of the first pair, are further concealed by lateral exten-
sions of the carapace, and by the long telson (Tel) bent forward
below them. The color is a uniform pale yellowish brown. When
buried in soft wet sand the crabs are thus well protected from their
granular environment; their only problem is that of respiration,
but the antennules projecting into the water above the sand together
form a respiratory tube.

When an Emerita is stretched out at full length (fig. 1 C) it is
seen to be a complete crab. A smooth, rounded, shell-like carapace
covers the gnathothoracic region of the body, a 6-segmented abdo-
men bears an extremely long, pointed telson (Tel), at the base of
which project the relatively large uropods (Urpd). On the under
surface, the region of the mouth and the feeding organs, and the
flagella of the second antennae, unless the latter are extended, are
completely covered by a pair of large valvelike plates, which are the
meropodites of the third maxillipeds (fig. 3 A, Mrpd). At the sides
of the latter are two elongate pterygostomial plates (fig. 1 B, Ptst)
flexibly attached to the edges of the carapace. The maxilliped plates
open ventrally like a pair of trap doors, and if they are lifted the
long plumose flagella of the second antennae will be seen looped
upon each other immediately above them (fig. 3 B). When the anten-
nae are artificially extended, the flagella project in curves outward
and posteriorly (fig. 1 A). Following the maxillipeds are five pairs
of legs, the first pair of which are directed forward against the
maxilliped plates, the second, third, and fourth pairs usually folded
under the carapace, and the slender fifth pair bent upward and for-
ward into the branchial chambers. The abdomen of the female has
three pairs of slender pleopods in addition to the uropods.

In its postembryonic development Emerita talpoida, according to
Smith (1877), goes through four zoea stages and a megalops stage.
The first zoea stage was not observed by Smith, but in the other
stages the zoea has a long rostral spine and two lateral spines on the
carapace, large eyes on short peduncles, and an abdomen of five
segments bent forward beneath the thorax. The telson is a broad
plate, with a wide, rounded distal margin. The mandibles in all the
zoea stages appear to be functional organs. In the second stage the

no. 8 SAND CRAB EMERITA TALPOIDA—-SNODGRASS 5

appendages include, besides the mandibles, two pairs of maxillae,
two pairs of maxillipeds, and the uropods. The third maxillipeds

Fic. 1.—External features of Emerita, Galathea, and Calappa.

A, Emerita talpoida (Say), female, with antennae extended, dorsal. B, same
with antennae concealed, left side. C, same, abdomen artificially extended, dor-
sal. D, same, abdomen detached, showing long apodemal plate of first tergum,
dorsal. E, same, cross section of body behind second legs. F, same, carapace,
pterygostomial plates, and branchiostegites, ventral. G, Galathea californiensis
Benedict, female. H, Calappa flammea (Herbst), carapace, ventral.

and the legs are still unsegmented rudiments. In the third zoea the
legs are better developed, and those of the fifth pair appear ; the gills
are now represented by slender processes above the first four legs.
In the fourth stage the full number of gills (nine on each side) is

6 SMITHSONIAN MISCELLANEOUS -COLLECTIONS VOL. I17

present, the uropods have two apical lamellae, but the telson preserves
the broad form of earlier stages. The megalops at once takes on in
many ways the form and habits of the adult. The second antennae,
Smith says, have assumed all the important features of the adult
antennae, including the mechanism for folding beneath the body,
but there are still fewer annulations in the flagella. The mandibles
have been reduced to the adult condition, the third maxillipeds form
large opercular plates, the abdomen and telson resemble these parts
in the adult. At Woods Hole the young Emeritas in the megalops
stage, according to Smith, may be taken at the surface of the water
in the towing net during late August and the early part of September.
Specimens in aquaria swim rapidly at the surface, but in a shallow
dish of sea water with sand on one side, they at once back themselves
into the sand “with an evident satisfaction and with an ease and
agility that could not have been excelled by their sand-loving parents
after months or years of practice.”

The first zoea stage of Emerita talpoida is said by Wharton (1942)
to lack the lateral spines of the carapace. The megalops, according
to Wharton, are amphibious like the adults, being well adapted for
both swimming and burrowing. In swimming, the megalops keeps
the abdomen extended, while the young adults swim with the abdo-
men flexed. The ventral pleopods of the megalops are used as swim-
merets, there being four pairs in the megalops stage, differing in
form from those of the adult.

It is customary to describe the body of a decapod crustacean as
divided into two parts, a cephalothorax covered by the carapace,
and a free abdomen, or pleon. However, there is a distinct head
structure in front of the carapace, or beneath its projecting anterior
end, which bears the eyestalks, the two pairs of antennae, and the
labrum. It is not intimately united with the body, except in Palinu-
ridae, and should be regarded as the true head of the animal. The
region covered partly or wholly by the carapace includes the segments
of the mandibles, the maxillae, the maxillipeds, and the legs, and is
hence really a gnathothoracic tagma, or section of the body. The
abdomen, then, constitutes a third part carrying the pleopods and the
telson. This concept of a triple division of the decapod will be followed
in the description of Emerita.

il. THE HEAD AND:ITS,.APPENDAGES

The head of most of the malacostracan Crustacea represents the
simplest type of head structure found among the arthropods, and
therefore, in an evolutionary sense, the first known stage of head

no. 8 SAND CRAB EMERITA TALPOIDA—-SNODGRASS 7

development. For this reason the writer (1951) has termed it the
protocephalon. A more complex secondary type of head structure
results from the addition of two or more postoral segments to the
protocephalon, as in the amphipods, isopods, myriapods, and insects,
in which the head bears, in addition to the eyes and antennae, the
appendages concerned with feeding.

The head.—The head of Emerita is a typical protocephalon ;
though it is not so well sclerotized or so strongly developed as that
of a crayfish or a lobster, it can be separated as a discrete head unit
from beneath the anterior end of the carapace (fig. 2A, C). On the
dorsal surface (A) is a V-shaped ocular plate (e) supporting the
eyestalks (ES), and behind it a larger, weakly sclerotized postocular
plate (m) flexibly connected with the base of the under lamella of
the projecting rostral part of the carapace. The small first antennae
(zAnt) arise anteriorly beneath the edge of the ocular plate, and at
the sides are the bases of the large second antennae (2Ant). On the
ventral surface (C), the head presents a broad calcified region, the
epistome (Epst), between and behind the bases of the antennules,
with lateral extensions to the bases of the antennae and irregular
postantennal wings (k). Supported on the posterior margin of the
epistome is the triangular labrum (Lm), which projects forward
inverted against the epistomal surface. If the labrum is turned
posteriorly (D, Lm) it will be seen to have covered a noncalcified
area of the epistome from which arise a pair of small setigerous
processes. In most decapods the postantennal wings of the epistome
(k, k) extend laterally to the bases of the inner lamellae of the
carapace folds, but in Emerita the intervening regions are not calcified.
At the basal angles of the labrum are the small mandibles (E, Md)
solidly united with the adjoining edges of the epistomal wings, and
behind the labrum is the mouth (Mth) flanked by a pair of small
paragnaths (Pgn). The mandibles and the paragnaths, however, do
not belong to the head.

The crustacean protocephalon is an anatomical fact, but its seg-
mental composition is an uncertainty. The origin of the second
antennae of the adult from the protocephalon might suggest that
the protocephalon contains the second antennal segment. The second
antennae, however, are postoral in their embryonic origin, and acquire
their definitive preoral position secondarily. The idea that the mouth
has migrated posteriorly from a more anterior position seems hardly
tenable if it is implied that it has pushed clear through one or more
segments. More probably, the ventral parts of the invaded segments
are simply folded forward around the sides of the mouth. That

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

something of this nature has taken place is indicated by the finding
of Tiegs (1940, p. 133), in his study of the development of a sym-
phylan, that much of the roof of the preoral cavity of the head is

Fear;
Ops

S cae, qo
eT “Dy ee

Ss
TMT
ee
Se
Qy
S
Pe ®.
3
WS

Ree

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sai

sy)
th

2 Sa se

>
5
a

4h uP Is
Ht
SSS

Fic. 2—EHmerita talpoida (Say), head, mandibles, and mouth region.

A, head (protocephalon) bearing eyestalks and first and second antennae,
dorsal. B, left eyestalk, and supporting plate of head (e), dorsal. C, head,
ventral, labrum normally turned forward beneath epistome. D, epistome, first
antennae, and bases of second antennae, ventral, labrum artificially turned pos-
teriorly. E, mouth region with associated labrum, mandibles, and paragnaths,
ventral. F, right mandible, dorsal.

derived from the premandibular (second antennal) segment. How-
ever, in the adult arthropod, no remnant of a postoral arc of even
the second antennal segment has been anatomically identified as such.
The identity of the dorsal are of this segment is equally elusive in
the adult. The origin of dorsal muscles of the second antennae in

no. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 9

the decapods on the anterior part of the carapace might suggest that
the rostral area of the carapace includes the dorsum of the second
antennal segment; yet in the adult anostracan branchiopod there is
clearly no segment between the protocephalic head and the man-
dibular segment. Recently it has been asserted (Henry, 1948) that
there is no second antennal segment. It would indeed simplify matters
very much to have this segment abolished, but unfortunately there
is too much evidence of its existence, at least in works on arthropod
embryology.

The eyestalks.—The slender eyestalks of Emerita (fig. 2B) are
3-segmented, with the smal! eyes on the ends of the long distal seg-
ments. The second segments are abruptly and flexibly elbowed on
the mostly membranous basal segments, which are borne on the ends
of the ocular plate (e) of the head, so that the eyestalks can be freely
protracted and retracted in a lengthwise direction.

The first antennae.—The relatively small first antennae, or anten-
nules (fig. 2 A, rAnt), project forward between the eyestalks. Each
antennule consists of a 3-segmented basal stalk, best seen from below
(C, D), and a pair of multiarticulate flagella, of which the ventral
one is more slender and a little shorter than the dorsal one. The large
basal segment of the stalk is of irregular form and articulates below
on the epistome ; the second segment bears ventrally a large setigerous
process. The flagella are fringed with short hairs and when the four
of them are held together they enclose a narrow space between them,
which is the usual exit canal of the respiratory system, but may
become inhalant when the crab is buried in the sand. The anten-
nular canal of Emerita, however, is not so well closed by setae as in
some other sand-burrowing crabs.

The second antennae.—The second antennae are organs of vital
importance to Emerita inasmuch as they are the food-collecting instru-
ments of these crabs. Each appendage (fig. 3 F) consists of a 2-seg-
mented basal stalk, of a middle part that contains two segments
which constitute the flexing mechanism, and of a distal part including
an elongate proximal segment, and the long, plumose terminal flagel-
lum. The two basal segments are evidently the coxopodite (Cxrpd)
and the basipodite (Bspd). The second is produced laterally into
a pointed lobe, but there is no true exopodite branch. The identity
of the other segments is difficult to determine. Schmidt (1915) in
his study of the antennal muscles of the crayfish accounts for seven
antennal segments, making the flagellum the dactylopodite. There
are, however, only five functional segments besides the flagellum

IO SMITHSONIAN MISCELLANEOUS: COLLECTIONS VOL. I17

in the antenna of Emerita, and the third and fourth are of irregular
form.

The third antennal segment (fig. 3 F) is largely membranous, but
its mesal part contains a strongly calcified convex plate supported
on the inner angle of the basipodite. The fourth segment presents
dorsally (B) a large, smooth, convex surface, but ventrally (F) it
is represented only by a Y-shaped bar that supports the fifth segment,
and articulates by each arm on the mesal plate of the third segment.
The two articular points lie in an oblique line (d-e) extending prox-
imally and laterally between the third and fourth segments, and it
is on this line as an axis that the distal part of the antenna is flexed
ventrally. The fourth segment, moreover, is braced laterally against
the basipodite by a short rod (f) in the ventral wall of the third
segment, which turns on the basipodite at a point in the axis of flexion.
When, therefore, the distal part of the antenna bearing the flagellum
is flexed ventrally on the oblique axis (d-e) between the third and
fourth segments, it not only turns downward and posteriorly, but it
goes over to the opposite side of the body, and the bar (f) braced
against the fourth segment prevents any other movement. The two
antennal flagella are thus assured of being symmetrically folded upon
each other (B); in most specimens the left flagellum is below the
right, but in some the right is below the left.

In the folded position of the antennae (fig. 3 B), the parts beyond
the lines of flexion are entirely concealed within the large meropodite
plates of the third maxillipeds (A, Mrpd), and the flagella are looped
upon each other. According to Weymouth and Richardson (1912),
Emerita analoga folds the antennae separately as the water runs off
the beach. The flagella are first allowed to trail out with the current,
and are then “folded up and withdrawn under cover of the third
maxillipeds with such a rapid motion as to escape analysis.” So
closely are the antennae shut in above the maxilliped plates, however,
that it would seem the plates must first open to receive them.

Inasmuch as the antennal flagella are specifically the food-collecting
organs of Emerita, the details of their structure become a matter
of special interest. Each flagellum of Emerita talpoida is cut into
about 150 small rings by circular joints that impart a flexibility to the
shaft as a whole. In cross section the shaft is rounded above and
somewhat flattened below (fig. 3 E). The plumose character of the
flagellum is due to the presence of eight rows of setae on the under
surface, there being duplicate sets of four on each ring, so that in
all there are about 1,200 setae on the entire flagellum. The setae
of the outermost rows (E, g) are long, tapering filaments flaring

no. 8 SAND CRAB EMERITA ‘TALPOIDA—SNODGRASS II
outward with incurved tips; the concave margins are densely fringed
with long slender hairs, giving these setae a featherlike structure.

The setae of the second rows (h) are somewhat curved outward ;

Ss
ne
YY

Dee ay

ZTE

GEE
4s
/;
Y/1

SSS.
Vp

SS
=
.

Fic. 3—Emerita talpoida (Say), second antennae and third maxillipeds.

A, ventral surface of anterior part of body, second antennae folded above
meropodite plates of third maxillipeds. B, second antennae in folded position,
ventral. C, a median process of antennal flagellum (£, 1). D, under surface
of a flagellar ring, showing position of setae. FE, cross section of flagellar ring,

with setae. F, proximal part of right second antenna, showing folding mecha-
nism, ventral.

their inner surfaces are armed with short, blunt, closely set processes
like the teeth of a comb. The setae of the third and fourth rows
(i, 7) are short and simple with fine hairs on their inner margins.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Finally, along the midline of the flagellar shaft, between the inner-
most setae, is a row of slender, tapering processes (J), all slightly
inclined toward the base of the flagellum. The tip of each of these
median processes (C) is obliquely truncate and appears to contain
a slight membranous depression. Weymouth and Richardson suggest
that these structures in E. analoga may be sensory organs. At D
of figure 3 are shown the relative positions of the several rows of
setae and the median processes on a single ring of the flagellum.

When the antennae of a dead specimen of Emerita talpoida are
extended (fig. 1A) they project forward from the body and the
flagella curve outward and posteriorly ; in this position the marginal
rows of setae are dorsal and ventral, with respect to the body. If,
however, the antennae are turned horizontally away from the ventral
surface as the crab stands vertically in the sand, or are carried into
this position by the downflow of the water on the beach, the antennae
will be rotated into a position in which the concave sides of the flagella
are away from the ocean, and the setae will now be spread out against
the descending current. Living specimens of £. analoga, buried in
the sand in the feeding position, are said by Weymouth and Richard-
son to hold the antennae “directed forward’? (presumably meaning
oceanward) and laterally with the tips curved outward. In this
position, these writers note, “the antennae thus present to the water
flowing against the concave side a very efficient straining apparatus.”
In other words, the opposite rows of setae are spread out as a fine-
meshed sieve against the downflowing water.

The stomach contents of Emerita analoga were found by Wey-
mouth and Richardson to be always of the same kind of material,
“chiefly shells of various diatoms, masses of brownish oily matter
apparently derived from the diatoms, radiolarians, foraminifera,
spicules of unrecognized origin, what were probably one-celled algae,
and considerable amounts of sand—about what would be obtained
by unselective straining of the water along the shore.” From experi-
mental feeding of E. analoga on suspensions of bacteria in liter jars
of sea water, Zobell and Feltham (1937-38) found that the crabs
can be sustained to a certain extent on bacteria alone, but that they
do not thrive on a pure bacterial diet as well as does the mussel M yti-
lus californianus, and that they are not as efficient as the mussel in
removing bacteria from the water. The antennae of Emerita in the
folded position above the plates of the third maxillipeds are separated
from the mouth by the second and first maxillipeds and the two
pairs of maxillae; the method by which food material is transferred
from the flagella to the mouth is, therefore, not open to observation.

no. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 13

Ill. THE GNATHOTHORAX

The gnathothoracic part of the body of Emerita is entirely covered
above and on the sides by the carapace (fig. 1 A, B). The anterior
dorsal margin of the carapace is produced into three small points
over the head, the posterior lateral areas are extended downward in
broad folds (B, bf) over the leg bases. In front of these folds on
each side is an elongate plate (Ptst) reaching from the base of the
second antenna to the base of the first leg, and flexibly attached to
the margin of the carapace. This plate is known as the pterygostomial
plate because it evidently represents the so-called pterygostomial
region of the carapace of an astacuran decapod. The posterior end
of the plate is not continued into the lateral fold (bf) of the carapace
behind it, but goes beneath the latter to be continued into a soft,
horizontal inner fold (F, Bstg) lying above the leg bases and closing
the gill chamber from below (E). This soft inner fold is the true
lower part of the branchiostegite (Bstg), the outer sclerotic fold
(bf) is a secondary extension of the branchiostegite beyond the base
of the inner fold. The branchiostegites themselves are folds of the
thoracic walls projecting from the sides of the back; their inner
lamellae are continuous dorsally with the mesal walls of the gill
chambers, which are the true lateral walls of the thorax on which
the legs are articulated (fig. 8B). In carcinology the inner wall of
the gill chamber is called either the epimeron or the pleuron, but for
general usage pleuron is preferable because the gill-chamber wall of
the crustacean evidently corresponds with the skeletal parts called
the pleura in other arthropods.

In order to establish a basis for a consistent nomenclature, it will
be of interest to follow the various modifications of the sclerotized
dorsal part of the body wall that occur among the arthropods. In a
simple, soft-bodied animal with ventrolateral rows of legs, such as
an onychophoran (fig. 4 A), the circumference of the body is divided
by the lines of the leg bases into a dorsum (D) above the appendages,
and a venter (V) between them. A simple sclerotization of the body
wall would then produce in each segment a dorsal plate, or tergum,
and a ventral plate, or sternum. If the tergum comes down to the
leg bases, the legs will be suspended from, or articulated on, its
lower margins, a condition found in some arachnids. More commonly,
however, as in the crustacean Anaspides (B), the dorsum of a tho-
racic segment contains a major tergal plate (T) covering the back,
and on each side a small pleural plate (Pl) carrying the outer artic-
ulation (a) of the leg. Usually, however, the contour of the segment

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Fic. 4.—Diagrammatic cross sections of arthropods, showing various types
of modification of the dorsal sclerotization.

A, dorsum entirely unsclerotized, as in an onychophoran. B, thorax of the
crustacean Anaspides, dorsal sclerotization differentiated into a principal back
plate, or tergum (T), and small laterotergal plates, or pleura (PI), carrying
lateral articulations of coxae. C, abdomen of an astacuran decapod, with small
tergal folds (tf) projecting over pleura. D, thorax of an astacuran decapod,
with tergal folds extended downward as gill covers, or branchiostegites. E,
thorax of a crab, Callinectes, with branchiostegites produced into lateral folds
(bf). F, thorax of a crab, Calappa, with branchiostegite folds depressed. G,
thorax of Emerita, with branchiostegite folds (bf) extended downward over
the legs, enclosing subbranchial canals (#). H, prothorax of an insect with
short lateral folds (paranotal lobes) of tergum. I, alate segment of an insect,
with tergal folds greatly extended horizontally and flexible at their bases,
forming wings.

No. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 15

is not so simple as in Anaspides. In the decapod abdomen (C) the
tergum is generally produced into short lateral folds (tf) overhanging
the pleura (P/). In the thorax (D) the pleura maintain a more or
less vertical position as the lateral walls of the body cavity, while
the tergal folds (#f) arise high up on the sides and descend as long
branchiostegites (Bstg) over the gills to the bases of the legs. In
most of the Brachyura (E, F) the outer walls of the branchiostegites
(Bstg) are extended laterally into sharp-edged folds, but finally, in
Emerita (G), the branchiostegite folds (bf) become thin ventral
extensions of the carapace. A lesser development of branchiostegite
folds is seen in the crab Calappa (fig. 1 H) in the form of flangelike
projections from the sides of the back, and in the anomuran Galathea
(G) there is a suggestion of the same thing in the presence of spiny
ridges (bf) above the branchiostegites.

In all the decapod forms above discussed, it will be seen that the
pleura form the true lateral walls of the thoracic segments, and carry
on their lower margins the lateral articulations of the legs. The
same is true for the thoracic segments of insects. In the insect
prothorax there are usually short tergal folds (fig. 4H, tf) over-
hanging the pleura; in the winged segments (1) the tergal folds are
greatly extended, flexible at their bases, and become organs of flight.
It is interesting to note, therefore, that the wings of an insect (I, W)
may be regarded as structures topographically homologous with the
branchiostegites of an astacuran decapod (D, Bsig).

The ventral surface of the gnathothorax of Emerita can be fully
seen only after removal of the appendages, so, unless specimens are
plentiful, it will be well to study first the mouth parts and legs,
described in the next section. On the exposed ventral surface (fig. 5)
there is to be noted an abrupt change in the sternal structure between
the region of the maxillae and maxillipeds (A) and that of the
pereiopods (B). The foramina of the legs (B, rL-5L) lie in regular
rows at the sides of the narrow median sterna (S). The third maxil-
lipeds (A, 3M xpd) arise lateral of the first legs (7), and the three
pairs of maxillipeds lie along oblique lines converging forward to-
ward the mouth; the second maxillae (2M) again have lateral
positions, and the first maxillae (1M) arise in membranous areas
at the sides of the mouth. The sternal sclerotization of the maxillo-
maxilliped region (A) is not segmentally divided; from the narrow
metastomal area it expands posteriorly into a large, diamond-shaped
plate between the bases of the maxillipeds in front of the first legs.
The maxillary region is extended on each side in a strong bridge
(mxB) to the base of the inner lamella, or doublure (Db/), of the

16 SMITHSONIAN MISCELLANEOUS. COLLECTIONS VOL, I17

pterygostomial plate of the carapace. The maxillary bridges evidently
represent the pleura of the second maxillary segment; they form the
anterior limits of the branchial chambers. It is to be noted that the
slender outer ends of the mandibles reach to the maxillary bridges,
on which in the Anomura generally the mandibles have their outer
articulations. The second maxillae have a lateral position because
of the respiratory function of the scaphognathites, and the first and
second maxillipeds are thrust forward between them, so that the
second maxillae are immediately followed by the third maxillipeds.

2Ant---

ro F i
AX 1L tMxpa_ 2Mxpa

Fic. 5—Emerita talpoida (Say), ventral skeleton of head and gnathothorax,
exposed by removal of appendages.

\
3Mxpd

A, ventral surface of head, mouth region, maxillary and maxilliped segments,
anterior to first legs (zl). B, ventral surface of pereiopod segments.

In the region of the pereiopods (fig. 5 B) the ventral sclerotiza-
tion at once assumes a regular pattern repeated in each of the five
leg-bearing segments. Between each pair of legs is a narrow sternal
plate or bar (S), which gives off laterally from its anterior end a
pair of slender precoxal sternopleural arms, and from its posterior
end a pair of similar postcoxal arms, which are continuous at their
outer ends with the pleura of the same segment. The adjacent sterno-
pleural arms of consecutive segments, however, are united to form
a series of interpedal brachia (ipB), in which the intersegmental
lines are marked by deep grooves.

The internal skeleton of Emerita consists of the usual inter-
segmental pleural and sternal apodemes of decapods. The sternal
apodemes of the thorax are four pairs of plates or arms diverging
forward and upward from the intersegmental grooves of the inter-

No. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 17

pedal brachia, the fourth pair arising from the brachia between the
fourth and fifth legs. The apodemes of the last two pairs are united
at their bases, the others are narrowly separated over the sternal
plates ; in no case are they bridged by expansions of their inner ends
as in the Astacura. A pair of divergent ridges in front of the first
legs are probably intersegmental between the third maxillipeds and
the first pereiopods. A pair of intermaxillary apodemes are united
in a transverse bridge, the so-called “head apodeme” behind the
mouth. The pleural apodemes are relatively small and are confined
to the thorax ; they arise on each side from the intersegmental grooves
of the pleuron (fig. 8B), and are Y-shaped, one arm joining the
corresponding sternal apodeme, the other going back to the one
following.

IV. THE GNATHOTHORACIC APPENDAGES

The segmental appendages of the gnathothorax are the mandibles,
the first maxillae (maxillulae), the second maxillae (maxillae), the
first, second, and third maxillipeds, and the five pairs of legs, or
pereiopods. The mandibles and the two pairs of maxillae may be
distinguished as the gnathal appendages; the maxillipeds are com-
monly regarded as thoracic, though in Emerita the first two pairs are
entirely concerned with the feeding function.

The mandibles.—The mandibles of Emerita talpoida (fig. 2 F)
are very small, though larger than in the other genera of Hippidae,
and are immovably united with the posterior edges of the epistome.
The basal part of each mandible (corpus mandibulae) is produced
mesally into a weak gnathal lobe (gnL), and bears a relatively large
2-segmented palpus (Plp). The two diminutive jaws lie entirely in
front of the mouth (E, Mth), with the gnathal lobes and the palpi
underlapping the base of the labrum (Lm). Though the mandibles
of Emerita clearly can have no use as biting or chewing organs, the
palps may have a sensory function, or, together with the gnathal
lobes, they perhaps serve to prevent the escape of food material from
in front of the mouth. A pair of small paragnaths (Pgn) underlie
the mandibular lobes.

In the zoea stages of E. talpoida the mandibles, as described by
Smith (1877), are evidently functional organs, since they have broad,
toothed gnathal lobes, though molar areas are absent. In the megalops
stage, however, Smith says, “the mandibles have become thin and
foliaceous and completely consolidated with the wall of the oral
opening.” A relatively large, indistinctly 2-segmented palpus is present,
but the gnathal lobe is a small thin flap as in the adult.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

The first maxillae—The small, flat first maxillae (fig. 6 A)
arise at the sides of the oral region and project mesally beneath the
mouth. The body of each appendage contains a small basal plate
and, proceeding from the latter, a pair of parallel bars, each of which
supports a large free mesal lobe, while at the base of the more distal
of these lobes there arises a small anterior lobe. The structure of
the first maxilla of Emerita is similar to that in other decapods,
and the small anterior lobe is commonly regarded as the endopodite,
according to which interpretation the other two lobes are coxal and
basal endites. It is of interest to note that the flat, weak, proximal
mesal lobe of the maxillula of Emerita is developed in other hippid
genera into a strong, toothed jaw (fig. 10 E).

The second maxillae.—These appendages (fig. 6 B, C) are larger
than the first maxillae, but they have essentially the same structure,
except for the presence of the huge scaphognathites (Scpg) attached
laterally on their bases. Three mesal lobes of each maxilla arise
from a common base and clearly correspond with the three lobes of
the maxillula (A). The maxillary scaphognathite is generally inter-
preted as the exopodite of the appendage, but its position suggests
that it might be a highly developed epipodite. The scaphognathites
are the active organs in the maintenance of water currents through
the gill chambers, as will be explained in the description of the
respiratory system.

The first maxillipeds.—The first maxillipeds (fig. 6D) are 2-
branched appendages borne on a pair of small basal plates situated
close to the midline of the body. The maxilliped branches are with-
out doubt true exopodites and endopodites. The endopodites of the
first maxillipeds are unsegmented; a sievelike arrangement of hairs
on their concave mesal borders lies below the mouth. The exopodites
are 2-segmented ; the apical segments bear brushes of long hairs and
converge against the sides of the labrum.

The second maxillipeds.——The maxillipeds of the second pair
(fig. 6 E) resemble those of the first pair in that each consists of an
endopodite and an exopodite supported on a small basal segment.
Their bases are farther apart than those of the first maxillipeds, which
latter in the figure (E) are seen between the basal segments of the
outer maxillipeds. The exopodites of the second maxillipeds bear each
an apical, brushlike segment similar to that of the first maxillipeds,
but the endopodites are 4-segmented, with the two slender distal
segments turned posteriorly. The long basal segments of the endopo-
dites are fringed mesally with wide, comblike rows of setae directed

no. 8 SAND CRAB EMERITA TALPOIDA—-SNODGRASS 19

1Ant

‘.

Fic. 6.—Emerita talpoida (Say), maxillae and maxillipeds.

A, left first maxilla, ventral (posterior). B, left second maxilla, flattened,
ventral. C, second maxiliae, with scaphognathites in natural position relative to
the labrum and bases of first antennae, ventral. D, first maxillipeds, ventral.
E, second maxillipeds, ventral. F, left third maxilliped, dorsal.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

toward each other, and laterally with long hairs that blend with large
setal brushes on the second segments.

The two pairs of maxillae and the first two pairs of maxillipeds lie
horizontally between the mouth and the infolded antennal flagella, and
the latter in turn are completely shut in ventrally by the plates of the
third maxillipeds. Weymouth and Richardson (1912) say that “the
mouth parts are admirably adapted for the manipulation of the minute
organisms which the antennae have strained from the waves.” Un-
doubtedly this is true, but it remains for some ingenious investigator
to discover just how the various mouth-part appendages perform
their function.

The third maxillipeds.—The third maxillipeds (fig. 6 F) have the
same segmentation as the legs; each consists of the endopodite alone,
there being no exopodite branch. The two appendages are widely
separated at their bases (fig. 3 A), but the broad meropodites (Mrpd)
come together mesally; the distal segments are turned posteriorly
(fig. 6 F) and are ordinarily concealed above the meropodites. A
short basal stalk of each appendage is formed by the coxopodite and
the basi-ischiopodite (fig. 6 F, Cxpd, BIscpd). The large meropodite
(Mrpd) is capped by a small carpopodite (Crppd), from which a
long slender distal arm of the appendage, composed of the propodite
(Propd) and the dactylopodite (Dactpd), turns posteriorly. The valve-
like meropodites are opened and closed by an axial rotation of the
appendages on their bases, but the mode of action by which they em-
brace and enclose the antennae has not been recorded. In the closed
position (fig. 3 A), fringes of long hairs on the proximal parts of
the appendages shut in the space behind the meropodites, covered
medially by the anterior end of the telson (Tel), which lodges the
posterior parts of the antennal loops (B). The narrow space between
the meropodites, and the triangular opening at their divergent anterior
ends are covered by overlapping setal fringes of the meropodites and
the distal segments (fig. 6 F). The outer edges of the meropodites lie
close against the pterygostomial plates (fig. 3 A, Ptst), and the large
first legs are pressed against their ventral surfaces. Whatever food
material the antennal flagella bring with them into the food chamber
above the meropodites is thus effectively imprisoned for transference
to the mouth.

The legs.—The legs of both the anomuran and the brachyuran
crabs have generally only six disinct segments, while the typical dec-
apod limb has seven, there being in the crabs only one apparent seg-
ment between the coxopodite and the meropodite. When a crab’s leg is
broken off, however, it leaves a small ring attached to the coxopodite,

no. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 21

which may be supposed to be a much reduced basipodite (fig. 7 B,
Bspd) detached from the ischiopodite. The functional second segment
of the leg, therefore, is termed the basi-ischiopodite (D, BIscpd).

The legs of Emerita, together with the uropods, are principally
organs for burrowing, though they must be used also for emergence
from the sand. The uropods serve both for burrowing and for swim-
ming. The burrowing activities of Emerita have been described by
several writers. Smith (1877) notes merely that E. talpoida burrows
backward by means of the thoracic legs and the uropods, while the
telson is kept appressed to the under surface of the body. Concerning
E. analoga, Mead (1917) says, “burrowing is accomplished by the
combined action of the uropods and pereiopods, the latter being the
more serviceable.” Animals with clipped uropods could burrow,
though more slowly than normally. MacGinitie (1938) gives only the
statement that the crabs bury themselves rapidly in the sand by use
of the anterior legs and the uropods. Weymouth and Richardson
(1912) record more detailed observations on the use of the different
appendages by E. analoga. The uropods, according to these writers,
appear to be the most important digging implements ; they strike up-
ward and forward in unison, thus thrusting the body downward and
backward if the sand is firm. The pereiopods take part in the burrow-
ing in different ways ; the second and third pairs thrust the body back-
ward, those of the fourth pair push outward, and the large first legs
are used to scull the body backward. “Under the combined action of
all these appendages the animal is carried diagonally downward and
backward until the anterior end of the carapace is just covered.”
According to Wharton (1942) E. talpoida, when preparing to bur-
row, backs up into the wet sand, and the uropods, rotating in unison,
throw the sand dorsally, while the body is forced backward. The first
legs work alternately pushing the sand laterally and anteriorly. The
fourth legs, and also the third and the second legs, Wharton says,
“move in unison laterally and posteriorly,” but the posterior move-
ment observed must be the relaxing stroke, since the statement follows
that these legs “move the sand anteriorly and force the posterior end
of the body into the sand.”

The first legs of Emerita are large, thick, and strong, with broad,
flat dactylopodites (fig. 7 A). As already noted, these legs project for-
ward and are closely applied against the meropodite plates of the third
maxillipeds, apparently serving to keep the plates tightly shut when
they enclose the antennae. The wide, broadly fringed dactylopodites,
however, adapt these legs also to their function in burrowing. The
other legs take a more transverse position and are ordinarily mostly

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

concealed beneath the branchiostegite folds of the carapace on the
sides and by the telson below (fig. 1 B). The second and third legs
(fig. 7 C, D) are similar in size and shape to each other. The dactylop-
odites have the form of short pruning hooks with broad bases and
decurved tips. Each dactylopodite is firmly hinged ventrally by a
double articulation on the end of the propodite, and when flexed up-
ward its wide base overlaps the posterior surface of the propodite.
Anteriorly the dactylopodite is braced against a strong projection from
the propodite (G, Propd). In either the flexed or the extended posi-
tion of the dactylopodite, therefore, the propodite and the dactylop-
odite of the second and third legs present a firm posterior surface for
pushing backward against the sand. On the other hand, a forward
push would appear to be more effective with the dactylopodites flexed
against the propodites. Evidently, then, the second and third legs
are constructed both for burrowing into the sand, and for emerging
from it. In the female the openings of the oviducts are on the coxae
of the third legs.

The smaller fourth legs (fig. 7 E) are turned posteriorly and up-
ward, and furthermore, are twisted in such a manner that the pos-
terior surfaces of the dactylopodites face outward. In Emerita tal-
poida the dactylopodites of these legs are simpler in shape than those
of the second and third legs, but they are likewise articulated ventrally
on the propodites and overlap the outer surfaces of the latter when
flexed. There is no bracing process of the protopodite against the
mesal surface of the dactylopodite, but the fourth legs, with the
dactylopodites in the flexed position, are clearly constructed to push
outward, as they are said to do by Weymouth and Richardson. In E£.
ialpoida there is little difference in the structure of the fourth legs be-
tween the female (EF) and the male (H).

According to MacGinitie (1938) the males of E. analoga “attach
to the female by the dactyls of their fourth legs, which are equipped
with a sort of sucker pad surrounded by stiff hairs. Sometimes they
hold on with both dactyls, at other times with only one. As viewed
under the microscope the sucking pad with its surrounding hairs does
not look very efficient, but the ease with which the males cling to a
female is surprising, for they have merely to place the dactyl against
any part of the carapace or appendage of the female to attain a hold.
However, the clinging is voluntary and not automatic.” A similar
habit has not been observed in the case of E. talpoida, but an examina-
tion of the end segments of a fourth leg of the male analoga reveals
a structure (fig. 7 1) quite different from that of talpoida (H). The

no. 8 SAND CRAB EMERITA TALPOIDA—-SNODGRASS 23

Fic. 7—Emerita talpoida (Say) and analoga (Stimpson), pereiopods
and uropod.

A, Emerita talpoida (Say), left first leg, ventrolateral. B, left leg of a crab,
Callinectes sapidus, showing narrow basipodite ring (Bspd) on which the rest
of the leg breaks off. C, Emerita talpoida, left second leg, posterior. D, same,
left third leg, posterior. E, same, left fourth leg, lateral. F, same, left fifth
leg of female, dorsal. G, same, anterior surface of distal segments of left second
leg, showing process of propodite against anterior surface of dactylopodite.
H, same, mesal surface of distal segments of fourth leg of male. I, Emerita
analoga (Stimpson), mesal surface of distal segments of fourth leg of male.
J, Emerita talpoida, posterior end of thorax of male, showing fourth and fifth
legs. K, same, right fifth leg of male, posterior. L, same, left uropod and
muscles from telson, in natural inverted position.

24. SMITHSONIAN MISCELLANEOUS: COLLECTIONS VOL. I17

dactylopodite of the fourth leg in the female of analoga is the same
as that of talpoida (FE), but in the male (1) the dactylopodite is broad
and hook-shaped as in the preceding legs; on its mesal surface is a
large oval depression, which is fully exposed when the dactylopodite
is extended, but it is overhung by a large brush of stiff hairs from the
propodite, and from its upper margin a brush of long hairs projects
beyond the tip of the dactylopodite, while a few delicate hairs arise
from its lower margin. Otherwise also the setal arrangement on both
the propodite and the dactylopodite is entirely unlike that of talpoida
(H). It seems clear, therefore, that the fourth legs of the male in
analoga are made for some special purpose, to which the correspond-
ing legs of talpoida evidently are not adapted, but it is difficult to
understand from the structure of the dactyls in analoga how this
special function can be adhesion. The means by which the male of
analoga holds the female with one or both of his fourth legs, there-
fore, needs closer observation on the living animals.

The fifth legs are small, slender, 7-segmented appendages (fig. 7 F)
that can be seen on an intact specimen only by depressing the telson
and straightening the abdomen. The coxopodites (J, 5L) arise close
behind the bases of the fourth legs (4L) ; the telopodites turn dorsally
and forward and are usually looped in the branchial or subbranchial
chambers (brC). These legs have diminutive apical chelae, the dacty-
lopodite being a small hook opposed by a short process from the prop-
odite. The distal segments are armed with long hairs, some of which
are directed proximally (F), forming brushes, suggestive that the
small fifth legs are used for cleaning the branchial chambers, or for
preventing the entrance of sand into their open posterior ends. In
the male (J, K) the coxopodites are particularly large and are pro-
duced distally into thick, soft papillae (Pen) on which open the genital
exit ducts.

V. THE RESPIRATORY. SYSTEM

The respiratory organs of Crustacea are primarily the gills, or
branchiae. When the gills are enclosed in branchial chambers, how-
ever, various accessories become necessary for efficient respiration:
there must be entrances into the chambers containing the gills, an
apparatus for maintaining currents of water through the chambers,
and an exit for the discharge of the respired water. In the decapods
all these parts together constitute the external respiratory system.

In Emerita the gills (fig. 1 E, Brn) are very closely shut within the
branchial chambers both laterally and ventrally by the branchio-
stegites (Bstg), and the posterior ends of the chambers are effec-

No. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 25

tively closed by the deflexed abdomen. The broad, descending lateral
folds of the branchiostegites (bf) enclose on each side a subbranchial
canal (4) between the inner fold of the branchiostegite and the legs.
Inasmuch as the inner folds taper in width posteriorly (F, Bstg), the
subbranchial canals and the branchial chambers become confluent in
the posterior part of the thorax. Anteriorly the soft inner folds of
the branchiostegites are continuous with the sclerotic pterygostomial
plates (F, Ptst), which are fully exposed on the sides of the body
(fig. 1 B, Ptst), except as they are partly overlapped by the first legs.
The lower margins of the pterygostomial plates lie close against the
bases of the first legs, so that in Emerita there are here no apparent
openings directly into the branchial chambers, as there are in some
crabs. At their posterior ends, however, the pterygostomial plates go
mesad of the lateral folds of the branchiostegites, where they become
continuous with the inner folds of the latter, and thus create openings
(fig. 1B, n) that lead directly into the subbranchial canals. The
pterygostomial apertures, therefore, would appear to offer a possible
way for the intake of respiratory water, which would have easy access
to the branchial chambers from the posterior ends of the subbranchial
canals, or, in the case of a reversal of the currents, the same openings
would serve as exits. This suggestion has only an anatomical basis,
and needs the support of experimental evidence.

Each branchial chamber (fig. 8 A, brC) opens anteriorly into a
pump chamber (PC) of the respiratory system, in which is lodged
the scaphognathite (Scpg) of the corresponding second maxilla. The
pump chambers are closed laterally by the pterygostomial plates. The
two scaphognathites converge anteriorly past the sides of the labrum
(fig. 6 C, Lm) into the triangular space below the bases of the anten-
nules(rAnt), which finally runs out into the median respiratory tube
formed by the close apposition of the four antennular flagella. If
water enters through the antennular tube, it will be split into two
streams by the apex of the labrum and thus drawn into the two pump
chambers ; or, if the current is reversed, the outgoing streams from
the pumps will unite at the bases of the antennules to be discharged
through the flagellar tube.

The vibratory movement of the scaphognathites is the force that
drives the water through the respiratory passages. In the decapods
generally the currents usually go from behind forward, but it has been
observed in some burrowing crabs and also in certain other species
that the currents can be reversed, presumably by a reversal in the
action of the scaphognathites, and the same is probably true of
Emerita. According to Smith (1877), when E. talpoida is buried,

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR) 117

the excurrent forms a small opening and a slightly boiling motion in
the sand. Mead (1917) says that by dropping small quantities of ink
into the water over a buried individual of E. analoga the current was
always found passing from posterior to anterior. On the other hand,
Weymouth and Richardson (1912), by the same experiment on E.

Ld ia

Fic. 8—Emerita talpoida (Say), respiratory organs and thoracic pleuron.

A, lateral view of thorax with pterygostomial plate removed and fold of
carapace cut away in the gill region, exposing anteriorly the pump chamber of
the respiratory system, with the scaphognathite in place, and posteriorly the
nine gills of the branchial chamber. B, left pleuron and leg bases of the gill-
bearing segments, gills removed but their positions shown. C, a single gill,
posterior. D, section of a gill, with paired lamellae on the axial shaft.

analoga, observed that the water is usually drawn into the antennular
tube when the animal is buried, though occasionally it goes in the op-
posite direction, but that when the crabs are resting on the surface of
the sand beneath the water or are swimming, the water is drawn in at
the sides and expelled through the antennular tube. Evidently, there-
fore, Emerita when buried adjusts the direction of its respiratory
currents according to conditions imposed by the sand.

no. 8 SAND CRAB EMERITA TALPOIDA—-SNODGRASS 27

There are nine gills in each gill chamber of Emerita talpoida (fig.
8 A, Brn). The first gill pertains to the segment of the third maxil-
lipeds (B, rgb), the others occur in pairs on the first four leg segments.
The gills all arise from large membranous areas above the bases of
the appendages. In the leg segments these areas lie within the pleura,
since they are closed below by marginal pleural bars bearing the coxal
articulations, so that the gills of the leg segments have the position of
pleurobranchiae. The membranous area containing the gill base on
the maxilliped segment (zgb), however, is not separated from the
articular membrane of the coxa, which fact gives this gill the status
of an arthrobranchia, though in position it falls in line with the other
gills, and evidently is serially homologous with them. The coxa of the
maxilliped is doubly articulated on the pleuron.

The gills themselves are crescent-shaped (fig. 8 C), and are closely
massed in the gill chambers (A, Brn) with their upper ends converg-
ing to a peak where the pleural wall is highest (B). Each gill consists
of two lateral rows of thin, closely set lamellae (D) arising from an
axial shaft on the concave inner margin. Below its middle the shaft
is connected with the pleuron by a short stalk.

VI. THE ABDOMEN, OR PLEON, AND ITS APPENDAGES

The abdomen of Emerita (fig. 1 C) consists of the usual six seg-
ments (1-6) and the telson (Tel) of the decapod abdomen. Dorsally
the tergum of the first segment (Zz) appears only as a narrow trans-
verse sclerite wedged between the thoracic carapace and the large ter-
gum of the second abdominal segment, but from its anterior margin
there is extended forward a large apodemal plate (D) inflected close
beneath the carapace, which gives attachment to numerous muscles.
The tergum of the second segment is as wide anteriorly as the cara-
pace, but the next three terga are successively narrower, so that the
margins of these four terga complete posteriorly the oval symmetry of
the body (A). The sixth tergum (C, 6) is a larger plate normally
having a ventral position (B, 6) when it is turned forward from the
fifth. The sixth segment carries the uropods (Urpd) and the telson
(fel):

The abdominal appendages of the Crustacea are appropriately called
pleopods, because the abdomen is known also as the pleon. Some
writers limit the term “pleopod” to the appendages of the first five
segments, and thus distinguish them from the uropods of the sixth
segment, but if the abdomen is the pleon, it is consistent that all its
appendages should be pleopods. The pleopods, however, are not

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 1i7

necessarily “swimming legs” as the name might seem to imply from
its derivation (Gr. pleo, sail, or swim), for some of them in the male
are usually genital accessories, and in the female egg-carrying organs,
the uropods alone being more consistently organs of aquatic locomo-
tion.

Fic. 9.—Emerita talpoida (Say), abdomen and pleopods.

A, left uropod, ventral. B, sixth abdominal tergum and right uropod, dorsal.
C, ventral surface of female abdomen and telson, with uropods in place, pleopods
of second, third, and fourth segments removed. D, second and third pleopods.
E, first pleopod with eggs.

In Emerita the adult female has three pairs of slender pleopods
pertaining to the second, third, and fourth abdominal segments (the
position of their bases is shown at C of figure 9). These pleopods of
the female are slender, 3-segmented appendages fringed with long
simple hairs (fig. 9 D), and are used as egg-carrying organs (E).
The first two pairs are much longer than the third. Corresponding
pleopods are absent in the male.

The uropods are present alike in both the female and the male. They
are relatively large appendages projecting from the sides of the sixth
abdominal segment (fig. 9 C, Urpd). Each consists of a long, thick
basal segment and of two flat, oval, closely appressed distal lobes with
apical brushes of long hairs. The uropods are freely movable append-
ages, since they have large, membranous basal supports on the body,

no. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 29

and are articulated only on pivotlike processes from the sides of the
tergum of the sixth segment (B). On the ventral side (A) a rigid
leverlike process projects from the base of the uropod into the artic-
ular membrane and gives attachment to two small muscles from the
sixth tergum, which evidently serve to turn the uropod outward on
the tergal pivot. The principal muscles of the uropods, however, arise
in the telson. A long outer muscle (fig. 7 L, mcl) runs the full length
of the telson to be attached on the posterior side of the base of the
uropod ; a much smaller muscle is attached anteriorly. The long mus-
cles evidently give a strong posterior stroke to the uropods, but when
the abdomen is flexed ventrally and the telson turned anteriorly, the
direction of the stroke will be forward. Emerita is said to swim always
backward, and in burrowing the uropods push forward.

VII. COMPARISON OF EMERITA WITH OTHER MEMBERS OF
THE HIPPIDAE AND WITH THE ALBUNEIDAE

Inasmuch as Emerita appears to stand alone as an antennal feeder,
it would be interesting to know the evolutionary steps by which it
became structurally adapted to its mode of gathering food, because
no halfway stage would seem to be practical. None of its relatives,
however, gives any helpful suggestion, since the other hippids have
gone their own way in the matter of feeding.

The genus Hippa in most respects is very similar to Emerita. Its
second antennae (fig. 10 B, 2Ant), however, do not appear to be in
any way adapted for food collecting ; though the large basal segments
are comparable with those of Emerita, the flagella are short, and the
whole appendages are brushlike by reason of their fringes of long
hairs, and are ordinarily folded beneath the bases of the antennules,
evidently to protect the respiratory passage between the latter. The
third maxillipeds of Hippa have broad meropodites similar in shape
to those of Emerita, but they are relatively smaller and only partly
cover the mouth parts. In both Hippa and Mastigocheirus the man-
dibles (B, Md) are even more reduced than in Emerita, but in these
two hippid genera substitute jaws have been developed from the mesal
lobes of the first maxillae. The first maxilla of Hippa cubensis (fig.
10 E) has essentially the same structure as that of Emerita (fig. 6 A),
but the proximal median lobe is a thick, strongly toothed jaw, those
of the two appendages closing together beneath the mouth. Each jaw
lobe is articulated on a basal rod of the appendage and has a second
articulation on the ventral skeleton by means of an articular arm of its
base. From the mesal surface (fig. 10 C) projects a strong process on

30 SMITHSONIAN MISCELLANEOUS -COLLECTIONS VOler Diy?

which is attached the tendon of a large adductor muscle (E, admcl).
Smith (1877) has described and figured the maxillula of Hippa
(Remipes) pacificus, which appears to be the same as that of H.
cubensis. For some reason, hard to guess, the hippids have all given
up their mandibles as feeding organs. Emerita has adopted plankton

LTO

ee
——

Cz
aun

5
ve

aun
CR
EQV R

]
ex

S
z=

HOS

Fic. 10.—Mastigocheirus and Hippa (Hippidae).

A, Mastigocheirus gracilis (Stimpson), ventral surface. B, Hippa cubensis
Saussure, head and mouth region, ventral. C, same, jaw lobe of first maxilla,
mesal. D, same, right first leg, ventral. E, same, left first maxilla, ventral.

feeding ; the food and feeding habits of Hippa and Mastigocheirus
appear to be unknown, but evidently their food is of such a nature that
a pair of jawlike organs was found necessary for its proper ingestion.

The second antennae of Mastigocheirus (fig. 10 A, 2Ant) are
relatively smaller than those of Hippa (B), but the first legs (A, rL)
have an extraordinary length, and the long, multiarticulate, brushlike
dactylopodites must have some special function when extended into
the water over a buried individual ; but if the function is food gather-
ing, it is not evident from the structure of these legs that the dactylop-

no. 8 SAND CRAB EMERITA TALPOIDA—SNODGRASS 31

odites could be brought back to the mouth. The corresponding legs
of Hippa (D) are large, but they reach only a little beyond the tips of
the antennules.

The antennules of the hippids are not so elaborately fringed as to
make a respiratory tube so well enclosed as that of the albuneids (fig.
11 A). In Emerita the four antennular rami are of about equal length
(fig. 2 A, D), in Hippa (fig. 10 B) the ventral rami are short, in
Mastigocheirus they are minute. The longer rami have fringes of
short hairs on their outer margins (fig. 10 B, rAnt) and a few very
small setae on their mesal margins. The space below and between the
antennular bases, however, is completely covered below by the large,
flat, brushlike apical segments of the exopodites of the first maxil-
lipeds, which are closely applied against the sides of the labrum, and
in turn are shut in below by the infolded second antennae. There is
thus formed at the base of the antennules a respiratory chamber which
is continuous distally with the interantennular tube, and divides prox-
imally into well-defined passages leading to (or from) the pump
chambers.

The members of the family Albuneidae, though clearly related to the
Hippidae, differ from the latter in various respects. The antennules
have each only a single flagellum, but the two flagella are held securely
together by fringes of interdigitating hairs along their dorsal and
ventral margins, forming thus an efficient respiratory tube, which in
Albunea (fig. 11 A) and Lepidopa is longer than the body. The sec-
ond antennae of these two genera are relatively short, but in Blepharip-
oda they are long and the flagella are plumose, resembling those of
Emerita. A distinctive feature of the albuneids, in contrast to the
hippids, however, is the presence of strongly developed mandibles of
the typical crab type of structure. The albuneids thus retain their
ancestral feeding organs. Benedict (1886) reports that the stomach of
a specimen of Lepidopa was found to contain setae of annelids, the
skin of a very small holothurian, and parts of the flagella of some
small crustacean.

The mandibles of Blepharipoda occidentalis converge anteriorly and
mesally behind the epistome, so that when closed the gnathal lobes
come together beneath the mouth. Each mandible (fig. 11 B) has an
elongate basal part expanded mesally, where it bears a broad, flat
gnathal lobe (gnL) and a 3-segmented palpus (P/p). The narrowed
lateral end is bent posteriorly, and from the point of angulation a
strong apodemal arm (Ap) projects laterally. The mandible is hinged
on the edge of the epistome, but has a specific mesal articulation in
front of the palpus, and a lateral articulation (a) with the ventral

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

skeleton at its recurved posterior end. The axis of rotation (a-c),
therefore, is oblique through the body of the jaw, and the apodeme
(Ap) stands almost at right angles to the axis. The apodeme supports
a large plate (mp) on which are attached antagonistic masses of mus-
cle fibers, and is thus an effective lever for abduction and adduction of
the gnathal lobe.

eZ
ly GE

Fic. 11.—Albunea and Blepharipoda (Aibuneidae).

A, Albunea oxyophthalma Leach, dorsal. B, Blepharipoda occidentalis Kan-
dall, right mandible, ventral. C, same, abdomen extended, dorsal. D, Albunea
oxyophthalma, left first leg, ventral. E, Blepharipoda occidentalis, distal seg-
ments of left first leg, lateral.

The third maxillipeds of the albuneids have exopodite branches, and
the meropodites are not widened. The first legs are strongly chelate
(fig. 11 D, E). The pterygostomial plates are continuous posteriorly
with the wide, soft branchiostegites, the two together forming on each
side of the thorax a broad fold flexibly attached on the edge of the
carapace, much the same as in Galathea (fig. 1 G). There are no outer
folds of the branchiostegites such as are characteristic of the Hippidae.
The albuneid telson is a short apical lobe of the abdomen (fig. 11 C,
Tel), which, when the abdomen is flexed ventrally, covers the bases
of only the fourth and fifth legs.

No. 8

SAND CRAB EMERITA TALPOIDA—-SNODGRASS 33

EXPLANATION OF LETTERING ON THE FIGURES

a, pleural articulation of coxopodite
(lateral articulation of mandible).

admd, adductor muscle.

An, anus.

1Ant, first antenna, antennule.

2Ant, second antenna, antenna.

b, sternal articulation of coxopodite.

BC, body cavity, haemocoele.

bf, fold of branchiostegite.

BIscpd, basi-ischiopodite.

brC, branchial chamber.

Brn, branchia, gill.

Bspd, basipodite.

Bstg, branchiostegite, gill cover.

c, epistomal articulation of mandible.

Cp, carapace.

Crppd, carpopodite.

Cxpd, coxopodite.

D, dorsum.

Dactpd, dactylopodite.

Dbl, doublure of carapace.

d-e, axis of ventral flexion of antennal
flagellum.

e, ocular plate of head.

Epst, epistome.

ES, eyestalk.

gb, gill base.

gnL, gnathal lobe of mandible.

ipB, interpedal brachium.

Iscpd, ischiopodite.

k, postantennal extension of epistome.

L, leg.

Lm, labrum.

mt, postocular area of head.

mcl, muscle.

Md, mandible.

Mrpd, meropodite.

Mth, mouth.

1Mx, first maxilla, maxillula.

2M, second maxilla, maxilla.

mxB, pleural bridge of maxillary seg-
ment.

iM «pd, first maxilliped.

2M-xpd, second maxilliped.

3M xpd, third maxilliped.

n, aperture into subbranchial canal.

PC, pump chamber of respiratory sys-
tem.

Pen, penis.

Pgn, paragnath.

Pl, pleuron.

Pip, palpus.

Propd, propodite.

Ptst, pterygostomial plate.

S, sternum.

Scpg, scaphognathite.

T, tergum.

Tel, telson.

tf, tergal fold.

Urpd, uropod.

V, venter.

W, wing.

x, subbranchial canal.

REFERENCES

Benenict, J. E.

1886. Revision of the Crustacea of the genus Lepidopa. Proc. U. S. Nat.
Mus., vol. 26, pp. 890-805, 8 figs.

Henry, Laura M.

1948. The nervous system and the segmentation of the head in the An-
nulata. Microentomology, vol. 13, pt. I, pp. 1-26, figs. 1-9.

MacGrnitig, G. E.

1938. Movements and mating habits of the sand crab, Emerita analoga.
Amer. Midl. Nat., vol. 19, pp. 471-481, 9 figs.

Mean, H. F.

1917. Notes on the natural history and behavior of Emerita analoga
(Stimpson). Univ. California Publ. Zool., vol. 16, No. 23, pp. 431-

438, 1 fig.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Scumipt, W.

1915. Die Muskulatur von Astacus fluviatilis (Potamobius astacus L.).

Zeitschr. wiss. Zool., vol. 113, pp. 165-251, 26 figs.
SmiTH, S. J.

1877. The early stages of Hippa talpoida, with a note on the structure of
the mandibles and maxillae in Hippa and Remipes. Trans. Con-
necticut Acad. Arts Sci. vol. 3, pp. 311-342, 4 pls.

Snopecrass, R. E.

1951. Comparative studies on the head of mandibulate arthropods. 118 pp.,

37 figs. Ithaca, N. Y.
Trecs, O. W.

1940. The embryology and affinities of the Symphyla, based on a study
of Hanseniella agilis. Quart. Journ. Micr. Sci., vol. 82, pt. 1, 225
pp., 41 figs., 9 pls.

WeymouTH, F. W., and RicHarpson, C. H.

1912. Observations on the habits of the crustacean Emerita analoga. Smith-

sonian Misc. Coll., vol. 59, No. 7, 13 pp., I pl.
Wuarton, G. W.

1942. A typical sand beach animal, the mole crab, Emerita talpoida (Say).
In Pearse, A. S., Humm, H. J., and Wharton, G. W., “Ecology
of Sand Beaches at Beaufort, North Carolina.” Ecol. Monogr.,
vol. 12, pp. 135-190, 24 figs.

ZoBELL, C. E., and FettHAmM, CATHARINE B.

1937-38. Bacteria as food for certain marine invertebrates. Journ. Marine

Res., vol. I, pp. 312-327.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 9

Roebling Hund

PRECIPITATION AND TEMPERATURE
IN WASHINGTON, D. C.,
FOR 1951 AND 1952

BY
Cc. G. ABBOT

Research Associate, Smithsonian Institution

(Pusiication 4087)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
MARCH 18, 1952

yLOID se

The Lord Baltimore Press

BALTIMORE, MD., U. & A.

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Roebling Fund

PRECIPITATION AND TEMPERATURE Py
WASHINGYON, DC} TOR 195T AND 1952
By C. G. ABBOT

Research Associate, Smithsonian Institution
A. PRECIPITATION AT WASHINGTON

Based on a cycle of 27.0074 days, I have predicted about 175 dates
each year when precipitation in Washington, expressed as the average
value per day, should exceed that quantity for all other dates. The

TABLE 1.—Statistics of Washington precipitation, 195I

(Values in inches)

Jan. Feb. Mar. Apr. May June
Average)! btds.) sence 0.053 0.152 0.169 0.154 0.183 0.277
perday J All other .... 0.085 0.040 0.040 O.III 0.031 0.337
Ratio are Eco cata 0.62 3.80 4.22 1.39 5.08 0.82
ALOtalappe. thse s «esis se'e 2.18 257, B17, 3.93 3.23 9.28
Normal pptak oe. oe.c =. 3.55 3.27 3.75 B27, 3.70 4.13
Percent of normal....... 0.61 0.79 0.85 1.20 0.87 2.25

July Aug. Sept. Oct. Nov. Dec. Year
Average. \Pidit sot. 8es% 0.028 0.088 0.089 0.036 0.1907 0.174 0.134
perday J All other .... 0.129 0.018 0.089 0.066 0.117 0.133 0.100
Ratio er Seeiorn ee hoe 0.22 4.89 1.00 0.55 1.68 1.31 1.34
Total pptieernck srieete ee 2.50 1.75 2.07 1.66 4.79 4.77. 42.50
INionmali ppt intaate jeuc tens 4.71 4.01 3.24 2.84 2137 3:32) Aes
Percent of normal....... 0.53 0.44 0.83 0.58 2.02 1.44 1.01

ratio of precipitation, Preferred dates, has exceeded unity each year
all other dates

for 18 consecutive years. The value of this ratio for the year 1951 is
134 percent. The expected value is 142 percent, and the average of it,
for 18 years, is 146 percent. Details for the year 1951 are given in
table 1.

Preferred days of 1951 had a higher average precipitation than all

1 See Smithsonian Misc. Coll., vol. 104, Nos. 3 and 5, 1944; vol. 111, No. 17,
1950; vol. 116, No. 4, 1951.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 9

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

other days in all months but January, June, July, September, and
October. In September the ratio was 1.00. In June 5.08 inches of
rain fell within 12 hours of preferred dates, so that the prediction on
these dates was but a half day amiss.

TaBLE 2.—Predicted dates when average daily precipitation should exceed
average daily precipitation for all other dates for the year 1952
in Washington, D. C.

“Preferred”
cycle places Jan. Feb. Mar. Apr. May June
Dae retohesenere 10 6 4, 31 27 24 20
Tse emt verte II Gi 5 28, 1 25 21
TT ire seereccrevanne 12 8 6 29, 2 26 22
[Vitaden eke 13 9 7 30, 3 oF, 23
Vi BEE ERs 14 10 8 4 28,1 24
Rigas tack 21 17 15 II 8 4
pa See eee 22 18 16 12 9 5
XW ies wisi 24 20 18 14 II 7
SKEW caaverstenste's 26 22 20 16 13 9
RAV ATTA Savers coneteters 27 23 21 V7, 14 10
B51 0 DS eer an ioe 31,4 27 25 21 18 14
EW tesa cors reine are 20, 2 25 22 18
DD QAI Damn Tamee 30, 3 26 23 19
“Preferred”
cycle places July Aug. Sept. Oct. Nov. Dec.
[eRe cicus te 17 13 9 6 2, 29 26
UT BAR Saye ee 18 14 10 7 3, 30 27
DUT Wee ce 19 15 II 8 4 28, I
TEV eghst sas Sterne 20 16 12 9 5 29, 2
Wiitetorgsseretteos 21 17 13 10 6 30, 3
XT Toa y otitis sieves 28, 1 24 20 17 13 10
OTT cesar sie's aes 29, 2 25 21 18 14 II
IRV of cs aysvayeasiets 31,4 27 23 20 16 13
PROVE Meter ctatets eve 6 20, 2 25 22 18 15
SOV ere eats avers Fi 30, 3 26 23 19 16
DON TUL Mr areter shercres II 4) 30, 3 27 23 20
A Ge es ae 15 II 7 31,4 27 24
RV TTS: 16 12 8 5 28, 1 25

Table 2 gives the dates in 1952 when the average daily precipitation
in Washington is expected to exceed the average daily precipitation
in this city for all other dates. Readers should neglect the first column
of the table, to avoid being confused. It merely gives, in Roman
numerals, the “preferred days” in the standard 27-day period. The
other columns show when these dates occur in all the months.

While it is expected that the dates given in table 2 for the months of
1952 will yield a higher average of daily precipitation than this aver-

NO. Q PRECIPITATION AND TEMPERATURE, WASHINGTON—ABBOT 3

age for all other dates of 1952 in Washington, the probability that any
individual date of the table will have any precipitation at all is not
greatly above 50-50.

The basic tabulation, on which table 2 rests, began with January
1924 and ended with December 1941. The length deduced for the pre-
cipitation cycle is 27.0074 days. In 379 cycles of this length there are
10,235.8046 days. From January 1, 1924, to December 31, 1951, there
are 10,227 days. Hence the 379th cycle begins on January Io, 1952,
as given in table 2.

This paper was prepared on January 14, 1952, but could not be
circulated in printed form until March 1952. Readers who may wish
to extend table 2 to cover the first three months of 1953 should note
that January dates are four days earlier than December dates, Feb-
ruary four days earlier than January and March and one day earlier
than February in 1953.

B. TEMPERATURE AT WASHINGTON

In previous papers ? I have noted a period of 6.6485 days in the
temperatures of Washington and New York, which has subsisted, on
the average, unaltered since 1910, though individual recurrences of it
are apt to depart one, two, and sometimes three days from regularity.

In table 3 I give the dates predicted for minima in 1951, the dates
when minima occurred, and the dates predicted for minima in 1952.
The predicted dates tabulated are those within a half day of those
accurately computed for minima, based on a cycle of 6.6485 days, with
zero date January 17.0000, 1946.

Figure 1 shows the relative frequency of departures of zero and
+1, 2, 3 days from the computed dates of minima of temperatures.

As another graphical example of the close accordance between
computed and observed minima of temperature in Washington I
submit figure 2. This gives the departures from normal temperatures
in Washington from March 21 to May Io, 1951. At the exact places
of minima, as computed with the cycle of 6.6485 days, from January
17.0000, 1946, heavy solid lines are drawn. As shown in the paper just
cited (Smithsonian Misc. Coll., vol. 111, No. 13, p. 6), a subsidiary
cycle of $6485 days was discovered. Dotted heavy lines represent the
places of minima of temperature at Washington for this subsidiary
cycle. It will be apparent that all the minima of the curve of tempera-
tures in figure 2 fall within one day of these heavy lines, indicating
places of computed minima.

2 Smithsonian Misc. Coll., vol. 107, No. 4, 1947; vol. 111, No. 6, 1949; vol. 111,
No. 13, 1949; vol. 116, No. 4, 1051.

TABLE 3.—Dates in 1951 when minima in Washington temperatures were

SMITHSONIAN MISCELLANEOUS COLLECTIONS

predicted and observed, and dates predicted in 1952

4
January
IQ5I predicted ...... 6 13 19 26
1951 observed ...... S) Tig BA} AID)
1952 predicted ...... 6 12 19 26
April
1951 predicted ...... 3) TOMA 23
1951 observed ...... 2 Ol 23
1952 predicted ...... TANS yids 27
July
1951 predicted ...... OMIZUIOY 25
1951 observed ...... 6 13 20 24
1952 predicted ...... 35-10) 16123
October
1951 predicted ...... 6 12 19 26
I95I observed ...... 10 20 29
1952 predicted ...... 4 Ir 18 24
15

30
28

February
8 15 22
8 15 24
8 15 21
May
13)-20:.27,
I2 19 29

Ir 18 24

August

8 14 21
Ouray 24)
I2"19 25

November
S seer
Seen 19

13 20 26

+15

28
28

March
7. TA 20) 27
10 22 26

612 919025

June
OmlOne2
5.10: 13. 10

Ft Qn Goi 127

September
alamo higwtez
8 18 24

Te Sutd yer

December
ATi rome!
10 16 22

3) or 16) 23

Fes

Fic. 1.—Relative frequency of minima of temperature at Washington

0 to +3 days from dates predicted.

VOL?) FIZ

29
29

31
27

NO. Q PRECIPITATION AND TEMPERATURE, WASHINGTON—ABBOT

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 10

Roebling Fund

PERIODIC TIES IN “THe
SOLAR -CONSTANT MEASURES

BY
C. G. ABBOT

Research Associate, Smithsonian Institution

(PuBLICATION 4088)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
MAY 28, 1952

TBe Lord Baltimore Press

BALTIMORE, MD., U. @ A.

Roebling Fund

PERIODICITINS IN Tin SOLAR-CONSTANT
MEASURES

By C. G. ABBOT!

Research Associate, Smithsonian Institution

INTRODUCTION

This paper, based on over 40 years of observations of solar radia-
tion, ties together the following conclusions:

The sun’s output of radiation varies.

It varies in at least 23 regular periodicities, all proceeding simultaneously.

The periods of solar variation are integral submultiples of 22% years.

Synthesis of curves representing the 23 periodicities reproduces the original

observations of the “solar constant” to within about 0.1 percent.

5. Synthesis of these curves for 12 years as a prediction, prior to the observa-
tions on which they depend, shows rough agreement with Mount Wilson
observations of the solar constant, in the years 1908 to 1920.

6. A much more satisfactory agreement is found between this predicted syn-
thetic solar-constant curve and the Mount Wilson determinations of the
march of contrast along the east-west diameter of the sun, of 1913 to 1920.

7. Higher contrast attends higher solar-constant values.

Gale LOE

In several former publications? I have discussed the periodic
changes in observed values of the solar constant of radiation.

For several years I have been investigating the effect on terrestrial
weather of these periodic changes in the sun’s emission. I had become
convinced by the earlier solar-constant studies, just cited, that the
sun’s radiation varies simultaneously in many regular periods, all

1] wish to express my sincere acknowledgments to L. B. Aldrich, Director
of the Astrophysical Observatory, who made the data available for this paper
and gave highly valuable criticisms; to Frederick E. Fowle, deceased, whose’
careful measurements of solar contrast appear in table 6; to Mrs. A. M. Bond,
deceased, whose critical judgment and accurate computations aided in the prepa-
ration of the data; to the many observers on high mountains in distant lands
who sacrificially kept up this long campaign of measurement; to Mrs. I. W.
Windom, who assisted in preparing this text; and to Miss M. A. Neill, who
continuously over many years greatly assisted me in keeping the observing
stations in operation.

2 Annals Astrophys: Obs., Smithsonian Inst., vol. 5, p. 250 et seq., 1932; vol. 6,
p. 178 et seq., 1942. Smithsonian Misc. Coll., vol. 111, No. 7, 1949.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 10

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

aliquot parts of 22$ years. I hoped, by using a long interval of scores
of years of an unbroken series of monthly weather records, that I
could discover from them all the submultiples of 223 years which yield
effective periodic variations of the solar radiation.

But I found that the variations of the atmospheric conditions from
time to time, some associated with the seasons and some with the
sunspot cycle, so badly confuse the phases of responses to solar varia-
tion that I could not be certain that all the suspected solar periodicities,
inferred from weather records, are real. Hence I felt constrained to
reinvestigate the observed fluctuations of the solar constant, to de-
termine directly which of the submultiples of 223 years are truly
periods in solar variation.

In former papers I have used 273 months as the master period, of
which the others are integral submultiples. My present work leads
me to prefer 272 months. All the periods which I have found lie
within less than I percent of being integral fractions of 272 months.

ADVANTAGES OF METHOD

Some investigators would prefer to submit the available solar-
constant data to a Fourier analysis based on 272 months. I prefer to
tabulate the data according to each suspected possible period. There
are several advantages in this method. In so doing, I divide the total
interval covered by the data into several parts, if periods are short
enough to furnish a large number of repetitions. In this way the
phases of features may be compared in the several independent tabu-
lations of one period. Graphs showing this procedure are given in
figure 1. Slight shifts,* from one to another of the successive tabula-
tions, indicate small corrections to the assumed period. The form of
the curve of fluctuation is determined by the tabulations. Also the
amplitude of the periodic variation is found. If it is too small to be
certainly exceeding the probable error, then the periodicity is to be
rejected altogether. Proceeding in this way, I found 23 periodicities
in solar-constant results which meet the tests of veridity just indicated.
Fifteen other periods were tabulated, but rejected. Each search in-
volved tabulating more than a thousand decade mean values of the
solar constant. The results appear in table 1.*

3 See the curves, 61/30, of figure 1, in comparison with table 1C, below.

4In tabulating any one periodicity, all the others exercise confusing influences,
which are not wholly eliminated, because of the small numbers of repetitive
columns going to make up the tables. Hence, irregularities in the curves of
figure I are caused by conflicting periodicities, in addition to the effects of acci-
dental errors of observation.

Talla) ‘Mal

uf mas

faye Baas a 2 CA peak ‘ae
9 Le, i kN fa Biss 4
oem ake Cate re *

Ne ‘ haat Oi 0 er ‘e = lah BS
ya (a aS ee ea nahh 7 oe ‘ ’
ean ye a aaa ri
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aoeds “SyJUOUI Ur payeoIpur a1e SpOliog “oS6I 0} OZ6I paAlasqo se uoleIpet
3 pue suoneuruejap yeyied sarynoesuoj—I “OI

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ATS AIM

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alt a
Re

NO. 10 SOLAR-CONSTANT PERIODICITIES—-ABBOT 3

It may aid to fix ideas on the method of tabulation to give an ex-
ample. Table 1C is a facsimile of the computation for the period
6 1/30 months. I select it as indicating how fractional parts of months
and of 10-day means are treated, so as to preserve the exact average
period. I had at first assumed that 61/15 months was the proper
length of period. The data were separated into three groups. The
assumed period corresponds with 18 1/5 10-day intervals. When the
mean values for the three groups were computed, they were plotted,
superposed. It was then apparent that the maximum ordinates shifted
progressively toward earlier dates, as time went on. This indicated
that the assumed period is too long by 4/700 of itself. Making this
correction, the true period is 6 1/30 months.

PREPARATION OF DATA

L. B. Aldrich, Director of the Astrophysical Observatory, and his
associates had painstakingly considered every circumstance affecting
every daily solar-constant observation, at all the Smithsonian mountain
stations in various lands. By consensus of three individual opinions,
they had assigned to every observed day its most probable solar-
constant value, as indicated by the checked results of all stations.
Many days were not observed at all. However, there was no decade
of any month, from 1920 to 1950, which did not have at least more
than one observation.

Mr. Aldrich having been good enough to place these daily solar-
constant results in my hands, I computed 10-day and monthly mean
values from them for the 31 years 1920 to 1950. To have them in
most convenient form for my use, I took their departures from the
value 1.900 calories per square centimeter per minute and divided
these departures by 1.940. Thus the results became expressed in
percentage departures of the solar constant from 1.900 calories. In
that form any well-evidenced periodic change resulting from a tabu-
lation shows at once its amplitude in percentage of the solar constant.
All values are positive as thus treated, which is convenient in tabula-
tion. These data are given in table 4, appendix I.

PERIODS FOUND AND NOT FOUND

With these clarifying remarks, I now introduce the results. The
following periodic changes in the solar constant were found well evi-
denced. Their approximate relation to 272 months and their ampli-
tudes in percentage of the solar constant are given in table 1A.

av rr if |

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4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

The following periodic changes, given in table 1B, if real, are too
small in percentage to be verified.

Tastes 1A, 1B.—Pertodicities in solar-constant observations

B. Periodicities sought

A. Periodicities confirmed * but not found

Period Amplitude Fraction Period Fraction
Months Percent of 272 Months of 272
27, 0.05 1/127 41/2 1/60
31/20 0.05 1/90 51/2 1/50
41/3 0.06 1/63 61/2 1/42
51/18 0.05 1/54 75/6 1/35
6 1/30 0.12 1/45 81/2 1/32
7 0.08 1/39 10 1/9 1/27
81/14 0.06 1/34 10 9/10 1/25
9 1/10 0.08 1/30 13 6/10 + 1/20
97/10 0.10 1/28 14 4/10 1/19

10 6/10 0.06 1/26 17 1/16

11 1/5 0.17 1/24 18 1/5 1/15

11.43 O.1I 1/24 19 1/2 1/14

12.0 0.20 wees 21 1/13

13 1/10 0.11 1/21 24 8/10 1/11

15 1/6 0.09 1/18 136 1/2

22 3/4 0.07 1/12

24 3/4 0.12 1/11

30 1/3 0.13 1/9

34 1/2 0.15 1/8

30 0.20 1/7

45 1/2 0.13 1/6

541/2t 0.13 1/5

68 0.25 1/4

OI 0.12 1/3

272 ann I

* The periodicities of 11.43, 12.0 (the periodicity of 12 months is not used in preparing
figure 4; if it were, that figure would present closer accord between the curves), and 24?
months were added to the list after search among the departures of the synthetic values, found
by summing 21 periodicities, from the observed solar-constant values. It is indeed curious to
find two periodicities both within 1 percent of 1/24 of 272 months. Both of them are ex-
cellently evidenced and of good amplitude. The 12-month period is of terrestrial, not solar,
causation. When one reflects that the pyrheliometer observes only about 70 percent of the
solar constant, the remaining 30 percent being supplied by our estimates of atmospheric
transmission, it is perhaps not surprising that the yearly (terrestrial) periodic error in the
solar-constant values is as large as 0.2 percent in amplitude. The periodicity of 243 months
was the only other one which could be discerned in a residual plot of differences, smoothed
by 7-month running means.

{ After this work was done, I computed a table of the periodicity 54 8/10 months in the
precipitation of Peoria, Ill., 1856 to 1939. It showed no periodicity of 548/10 months, but
four strong, well-shaped periodicities of 54 8/10 + 4=13 7/10 months. Hence I think the
sun’s radiation has a periodic variation of one-twentieth of 223 years, though it did not im-
press me as real in the tabulation of the solar constant.

All periods of these two lists were separately sought for by tabulat-
ing over 1,000 solar-constant 10-day means for each suspected perio-
dicity. The investigation does not cover entirely the years 1922 and

NO. I0 SOLAR-CONSTANT PERIODICITIES—ABBOT 5

1923. I have elsewhere discussed the large solar change observed in
those years.® I still think it was a real one. But it may be either a very
unusual sporadic solar change, or it may be a periodic change related
to a longer period than 272 months.

CONCERNING DOUBTS OF SOLAR VARIATION

For those who do not have intimate association with the Smith-
sonian observations of the solar constant of radiation, it seems diffi-
cult to accept the results as having the high degree of accuracy claimed
for them. Observers, familiar with the clouds, dust, and water-vapor
load which the lower atmosphere bears to make it milky, do not readily
visualize a sky so clear that, if one holds his little finger at arm’s length
before the sun, the sky seems deep blue right down to the sun’s edge.
But even if the superior excellence of stations like Montezuma, Table
Mountain, and St. Katherine be granted, it still seems incredible to
many that the fraction, amounting to about 30 percent of the solar
constant, cut off by the atmosphere, can be so correctly estimated that
variations of the order of 1/10 percent of the solar constant can be
evaluated.

Still more doubtful does it appear to many that, lacking any theo-
retical support, it can be proved from the observations that the solar
variation consists of 23 simultaneously operating regular periodicities,
all aliquot parts of 22% years. Yet it seems to me this cannot longer
be doubted. I have tried to demonstrate by a couple of examples that
it is necessary to use integral fractions of 223 years, rather than any
other intervals, to represent the the sun’s periodic variation. The two
periods I have chosen to experiment upon are those which are 1/7 and
1/45 of 22? months. In figure 1 the longer period is plotted as 39
months.

I made a new tabulation in four parts for a period lying between
1/45 and 1/44 of 223 years. It was assumed to be 6§ months, or
Ig 10-day intervals. In each of the four groups tabulated there are
14. columns. Taking the mean values, they are as plotted in figure 2,A.
Evidently, if the four mean results were combined directly, they
would so contradict each other that the general mean would show
no periodicity at all. But the principal feature, marked A at its right-
hand edge in each plot, is equally displaced from curve to curve to-
ward the left by about 6 10-day intervals. The displacement is 19

5 Monthly Weather Rev., U. S. Weather Bureau, February 1923. Proc. Nat.
Acad. Sci., vol. 9, No. 6, pp. 194-198, 1923. Smithsonian Misc. Coll. vol. 77,
No. 5, 1925 (see fig. 11); vol. 80, No. 2, 1927.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

10-day intervals, in all, from curve I to curve IV. Between these
curves I and IV lies a stretch of time of about 800 Io-day intervals.
Hence the period should have been taken less than 6$ months by
19/800 xX 64=0.146. Subtracting from 6.163, this yields a corrected
period of 6.017 months. Within the error of determination, this checks

Fic. 2.—The periodicity 6.033 months, confirmed by the displacement of the
feature A gradually from I to IV, when the period is assumed to be 64 months,
as shown in figure A. In figure B this displacement is adjusted to a period of
6.017 months, which nearly agrees with the true period, 6.033 months.

with 6.003, which is the period given in table 1C. Having displaced
curves II, III, and IV by 6, 12, and 19 10-day intervals respectively,
and having taken the general mean of the four and plotted it, the
result appears in figure 2,B. It is to be compared with the curve of
6.033 months above it, representing the mean value as given in
table 1C. It must be admitted that the agreement is striking.
Proceeding similarly, I computed two curves ® for the seventh of

6 There being but four columns in these part computations for 39 and 37
months, the plots of the results are very ragged, owing to the disturbing in-
fluences of 22 other periodic factors superposed.

ABBOT

SOLAR-CONSTANT PERIODICITII

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nmoisn(niwy

HANMTPNONnDA

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. (EL7

223 years, assumed as 39 months. In this new tabulation I used
monthly mean values, instead of 10-day means, as had been done in
computing for the curve shown in figure 1. I also computed two
curves for a period of 37 months. They show opposition rather than
similarity. It now appeared that in both the 39-month and the 37-
month computations, the principal features were displaced toward the
right in the second half of the 31-year interval. The corrected interval
from the 39-month tabulation is 394 months. Plots of the 37-month
tabulation shown in figure 3,A indicated a displacement toward the
right of 8 months in an interval of 180 months of time. This gives

a positive correction of Sx 37=1.6 months. Thus combined, the
180

contrary curves of figure 3,A yield the lower curve of figure 3,B.
Thus the 37-month tabulation yields an adjusted period of 39.6
months, closely agreeing with that yielded by the adjusted 39-month
tabulation which was 39.5 months. This later period agrees within

slightly more than I percent of being mE , Or 39.0 months. (See

figure 3,B.)

If critics feel that still more evidence is needed to prove that only
integral fractions of 223 years are to be found in the solar variation,
I will remind them that many of the periodicities plotted in figure I
show integral fractions of the periods in question superposed upon
them. Conspicuous examples in figure I are periodicities of 154, 344,
39, 454, and 544 months.

ACCURACY OF DATA

As shown in Annals of the Astrophysical Observatory of the Smith-
sonian Institution (vol. 6, p. 163), the comparison of daily solar-
constant values, independently measured at stations thousands of
miles apart, in opposite hemispheres of the earth, extending over
many years, yields a probable error for a well-observed solar-constant
0.164

value, resulting from work of two stations on a single day, of

V2
percent or $ percent. Using the familiar relation (the probable error
of a mean is that of the individual divided by the square root of the
number of values), this indicates that a 10-day mean of good quality
should be assigned a probable error of 1/25 percent. Then if nine
such 10-day means are tabulated in searching for a solar periodicity,
the probable error of their mean becomes only 1/75 percent. These
considerations indicate not only that real solar variations of 1/10
percent of the solar constant might be detected, but that the features

NO. IO SOLAR-CONSTANT PERIODICITIES—ABBOT 9

of the march of a periodic variation of this small amplitude would
appear well delineated from a tabulation.

To be sure, these optimum conditions do not always prevail. Not
infrequently no more than three or five days of a decade yielded solar-
constant observations. Often no more than one station reported. Dur-

ASS Pe oo
Wek MT AAA |
NYCI [Vet la
A, | P| TY
Te ee
Meetect tach Pk “Ios

A B

Fic. 3.—The periodicity of approximately 7X 272 months, tested just as the
periodicity of approximately 1/45 < 272 months was tested in figure 2.

240

230

220

240 220

230

ing parts of the year less favorable conditions prevailed at one or
other of the stations. Such is the case at Table Mountain from March
through June, and at Montezuma from November through January.
(See figs. 7, 8, pp. 70, 71, Annals, vol. 5.)

On these accounts it need not surprise us that, as shown below,
while the sum of periodic variations represents the variation of
monthly mean solar-constant results to within an average deviation of
1/10 percent, much larger departures sometimes occur. However,
divergences depend not only on accidental ertors of the observations,
but, in part also on imperfect determination of the form, amplitude,
and period of the periodicities, for reasons explained above.

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

SUPPORTING EVIDENCES OF VERIDITY OF PERIODICITIES

There are several indications, not flowing from a consideration of
probable errors, that strongly support the veridity of periodicities
here disclosed:

I. In tabulating periodicities, the data have been treated inde-
pendently in several parts. That is to say, there being nearly 1,100
consecutive 10-day means covering an interval of 30 years, it is pos-
sible to tabulate in three or more groups, each with numerous columns,
all periodicities of less than 20 months in length. For periodicities of
between 20 and 40 months I use two tables, covering consecutive
intervals of time. (See fig. 1.) Unless these independent part-tabula-
tions agree within their measure of accuracy to indicate continuance
of the same form of periodic variations, and with maxima in the
same phase throughout the whole time, then such a supposed period
is thrown out as nonexisting. For periods exceeding 40 months, the
data were not numerous enough to be thus separated into several
groups.

2. There is an integral relationship between the periods disclosed.
All the periods, which the first criterion certifies as veridical, are, to
within a deviation of 1 percent, integral submultiples of 272 months.
For example, those approximately 91, 68, 54, 45, 39, 34, 30, and a
dozen others of shorter period, are all integral fractions, to within
I percent, of 272 months. We know that a period of about 272 months
is related to the average sunspot period of 114 years, and it was found
by G. E. Hale in the behavior of sunspots and magnetism. It is also
approximately the period discovered by meteorologists in many cli-
matic phenomena, as well as by Douglass in the growth of trees.

I cannot but think that the fact of the integral relationship, each
to each, of the solar-radiation periodicities here disclosed, and the
relationship of all of them to a master period of 272 months, well
known in other solar and terrestrial phenomena, strengthens the case
for validity of these periodicities. If that be granted, surely the
existence of these integral solar-radiation relationships, so reminiscent
of the overtones of the vibrations of musical instruments, is a phe-
nomenon well worth investigating by astronomers and by students of
hydrodynamics.

I have just stated three arguments for the reality of numerous
regularly periodic variations of the output of radiation from the sun
as follows: A. Measurements whose small probable error is con-
sistent with the amplitudes of the apparent periodicities display them.
B. Tabulations of a chosen periodicity, with the data separated into

NO. IO SOLAR-CONSTANT PERIODICITIES—ABBOT II

independent groups, covering successive time intervals, show sepa-
rately the periodicity in similar amplitudes, forms, and phases. C. The
periods are integrally related, each to each, and all are approximately
exact integral submultiples of 272 months, itself a well-known period
in other solar and terrestrial phenomena. A fourth supporting evi-
dence is to be referred to later.

The argument B is undoubtedly the most telling. In order to display
its full weight, I give, in figure 1, a résumé of all the periodicities
which I consider real. It is my firm expectation that scientists who
examine without bias the arguments A, B, and C and carefully scan
figure 1 and table 1C, will yield to the conviction that the sun’s con-
tribution of radiation that warms the earth varies in a complex way.
In short, they will admit that, like the overtones of a musical note,
the radiation of the sun varies simultaneously in a period of approxi-
mately 272 months, and in periods, exceeding 20 in number, which
are integral submultiples of approximately 272 months. If scientists
go thus far, I cannot but think they will go farther and investigate
theoretically the hydrodynamics of the phenomenon.

PERIODICITIES OF 22% AND 113 YEARS

I have not tabulated the data so as to display the periodicity of
272 months, because the values are insufficient. There would be too
few repetitions to fairly fix the form of this curve. As for the perio-

dicity of 222 = 36 months, though it is the well-known 114-year

sunspot period, it is inconspicuous in the variation of the solar
constant. I have twice sought for it. First, I tabulated the original
data in columns of 136 months and smoothed their mean values.
Second, I smoothed by 7-month running means the residual depar-
tures, which separate the original data from the synthetic reproduction
of them in figure 4 by 23 periodic terms. Neither treatment gave con-
clusively a periodicity of 136 months. Its well-evidenced weather in-
fluence, I think, is attributable to fluctuation of the intensity of the
bombardment of the atmosphere by electric ions, acting as centers of
condensation of water vapor and dust, as sunspot numbers wax and
wane.

GRAPHS OF RESULTS

Figure 1 is introduced to emphasize the force of the argument B
by a graphical appeal to the eye. The figure shows the mean result of
every partial tabulation of the values used to compute table 1A, and
also the general mean of these partial tabulations for almost all perio-

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

dicities included in table 1A. Curves for periodicities of 21/7 and
31/20 months are given on a scale of abscissae 24 times as great
as the other curves. Horizontal lines in figure I are separated by
1/10 percent of the solar constant. The curves for periodicity 2 1/7
months are given on a scale of ordinates twice as great as that used
for all others. Up to a periodic length of 223 months, all the curves
are plotted at 10-day intervals. Periodicities of 22% months and
longer are plotted in monthly intervals. Of periodicities less than
223 months in length, one, that of 9 1/10-months period, is shown
smoothed throughout by 5-decade running means. It has a small
amplitude and would perhaps have seemed doubtful to many had
not running means of 5-decade values been shown, instead of the
separate 10-day mean values. This smoothing brings out plainly the
similarity of the partial tabulations.

The amplitudes of the 23 periodicities plotted in figure 1 may seem
to some critics too small to be of any significance. Not so. For it is
shown in figure 4 that the synthesis of these 23 periodic fluctuations
produces a curve closely matching, and of the same amplitude of
variation as, the curve of original observation. A 12-month period
of terrestrial origin with amplitude of 0.2 percent is not introduced
into figure 4. Its inclusion would improve the agreement there. No
additional regular periodicities were discernible. The analysis appears
to be exhaustive.

As the periods grow longer, they are apt to display integral sub-
multiples riding upon the period under examination. This is strongly
marked with the period of 154 months. It shows seven subperiods of
21/7 months very plainly. Similarly the 304-month curve shows also
the 6 1/30-month influence. The 344-month curve shows influence of
the 114-month period. Other examples are obvious. Note the curves
for periodicities of 543, 68, and 91 months shown in figure 1. Owing
to superposed periods of less length, these long periodicities had to
be smoothed by 5- or 7-month running means.

In addition to the direct mean results for each period, I give in a
few cases also the smoothed mean, resulting from taking 5-value or
7-value running means for the entire length of the periodicity under
consideration. These smooth curves give a more convincing and truer
idea of the periodicities, thought to be real, than do the rougher direct
means, affected by accidental errors of observation and influences of
extraneous periods. Readers should bear in mind that the knicks in
the broken lines, which look so large, really average less than 1/10
percent of the solar constant. This bears witness to the high accuracy

NO. I0 SOLAR-CONSTANT PERIODICITIES—ABBOT 13

of the Smithsonian solar-constant observing. Its probable error has
been discussed above.

INTEGRAL RELATIONSHIPS

I had long been of the opinion that the regular periodicities of solar
variation are all integrally related to approximately 272 months. This
impression is supported by the fact, so obvious in figure 1, that the
longer periods shown, themselves being integrally related to 272
months, have in several instances shorter periodicities riding on their
backs, which are integral submultiples of them. Further proof of the
integral relationships is shown in figures 2 and 3, already described.

Assuming that this integral relationship to 272 months is a condi-
tion necessary to the real existence of a regular period in solar varia-
tion, the number of such periods that are of considerable amplitudes
seems not to exceed 23. At least a rather extensive search has not
yielded others strong enough to be certainly real. If these be all, and
their forms and amplitudes are as shown in figure 1, then a synthesis
of them ought to represent the march of solar variation from 1920 to
1950, except for the interval of 1922 and 1923, when exceptionally
large solar variations were observed and which is excluded from this
analysis. I have made such a synthesis, and compare it with the
march of the solar variation in figure 4.

SYNTHESIS OF PERIODICITIES

To determine the quantities plotted in figure 4, I have computed
the departures, plus and minus, from the mean ordinate for each
smoothed periodicity, as expressed monthly, which together fix the
form of its curve. This gives, in each case, a short series of small
monthly departures suitable to the form of each periodicity. All the
tabulations begin with August 1920 as zero time. In table 2 they are
all tabulated in the smoothed form actually used in preparing the
synthetic curve shown in figure 4. In computing the mean periodic
forms, and afterward in using them for synthesizing the solar-constant
values, I allow for fractions of a decade, or of a month, by adding or
withdrawing a value from certain columns, or at appropriate intervals
in synthesizing, so as to preserve the correct period.

I tabulate these series, end to end, over the whole interval of more
than 30 years. Thus I make a great table of 23 columns and 367 lines.
Adding algebraically the plus and minus values of the lines across
the table, I find the total synthesized monthly departures, in ten-

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

thousandths of the solar constant, from the mean solar constant 1.94
calories. The results, covering 367 months, are compared in figure 4
with the monthly observational values recorded in table 4.

CLOSE AGREEMENT BETWEEN SYNTHESIS AND OBSERVATION

Table 3, below, shows the high degree of accuracy with which
the synthesis of the original 21 periodicities (before those of 11.43
and 243 months were found) corresponded to the observations.

These results came from the comparison of observation with the
synthesis of 21 periodicities. The average departures are reduced
below these figures when periodicities of 11.43, 12.0,7 and 243 months
are introduced. The value for the best 233 months then becomes 1.00-
tenths percent. The larger average departures prior to July 1926 are
attributable to the then imperfect development of the “short method”
of solar-constant work. The larger departures after 1945 are thought
by Mr. Aldrich to be caused by temporary errors in the scales of
pyrheliometers used in the field. He hopes to correct this discrepancy.

Some minds may still prefer to think that the solar-constant ob-
servations do not prove the variability of solar radiation. They may
point out that the average deviation of the observations from their
mean is 0.15 percent, and the average deviation of the synthetic curve
from that of observation is still 0.10 percent. They may urge that this
amount of improvement is not sufficient to warrant belief in the thesis
that the sun’s radiation varies in the discovered 23 regular periods,
all integral submultiples of 272 months.

Such critics may be reminded that the “weight” of any measure-
ment, that is, its claim to respectful recognition, is proportional to the
number of observations that enter into the result; but the probable
error (proportional to the average deviation from the mean) is pro-
portional to the square root of the number of observations. It follows
that the ‘‘weight,” or credibility of a solution, is proportional to the
square of the average deviation of its components. Hence the weight

of the solution here advocated is (25) =2.25 times the weight of

the conclusion of an invariable sun.

But it must also be considered that a certain irreducible minimum
of accidental error, comparable in a graph to the teeth of a saw, ad-
heres to the solar-constant observations. Whatever excursions from
the mean value may be produced by real solar variations, these acci-

7 The 12-month period is not used in preparing figure 4; its use would improve
the agreement of the curves.

TABLE 2.—Twenty-three solar periodicities in ten-thousandths of the solar
constant, based on August 1920. Also the 12-month terrestrial

period, same umt

21/7 M: +2 —2. 31/20M: o —2 +2. 41/3 M: —1 —2 +3 +0.
51/18M: —1 +o —2 +2 +2. 61/30 M: —4 —1 +3 +6 +0 —5.
7M: —1 +1 +5 +2 —1 —1 —2. 81/14 M: —2 —2 —1 —1 41 +1
+3 +2.
o1/1oM: —2 —4 —3 —1I +o +2 +3 41. +o.
97/1oM: —4 —3 —I +1 +5 +5 +2 —1 —4 —3.
106/10M: —1I —1I —I —1 —3 +1 +1 +2 +3 +1 —1.
11/5 M: —4 —2 +o +3 +1 +9 +3 —1 +4 —2 —8.
Bigg chy ai O tl 3) 4d 3 348
131/1oM: +1 +4 +3 —2 —6 —4 +2 +2 +1 +o —2 41 443.
1351/6 M: —3 —6 —6 —1 +0 +2 +1 +2 +3 +2 +o +o +2 +1 +41.
223/4 M: —1 +1 +o 41 +1 +1 +1 +1 Ho +o +1 +2 +3 +3 +2 +2
Se ah 2 S33) 21.
2a AM ee 2) 3) 3-4 4-4 3 3: 2 ft 02
—5 —7 —2 =o +o +o +o —I —1.
301/3 M: +6 +5 +4 +3 +3 +4 +3 +1 +1 =o to So —1 —3 —5 —6
—6 —5 —5 —6 —6 —4 —3 —2 —1 —1 +0 +3 +3 +4.
edie ar Oa ee eo Oat de eee O
4-8-7 4-004 S20 =) 20 Pt F243 4-3 i-4 eS tS) Fert
—I —3.
39M: —4 +1 +2 +2 +2 +2 +1 +1 +1 +2 42 +4 +6 +8 +10
palOc 8 cd wld (eeBeh4 “RS aS. cba: 3 a3) ced sek apg 6
—8 —I10 —10 —I0 —9 —9 —9 —8 —6.
4590/2 Ms 53 S43) 3.272 ST 0) EE SEE 4-3-4 6 16 3. be) x1
SO) 8 8 ae SE he he che 0 ks) es
—4 —3 —2 +o +1 +1 +o —2 —3 —4 —2 —2 —1.
541/2 M: +4 +4 +5 +6 +6 +7 +7 +7 +7 +6 +6 +6 +5 +3 =o —1
—I —I —2 —4 —4 —3 —3 —2 —2 —2 —2 —3 —2 —3 —2 —4
—5 —4 —3 —4 —2 —3 —3 —4 —3 —2 —I —I —I —2 —1 +0
+o —2 —2 —1 +1 +2.
68 M: —7 —5 —4 —4 —4 —6 —6 —8 —I2 —13 —I2 —9 —5 —4 —2
—3 —2 —2 —8 —II —II —I10 —6 —6 —4 —3 —4 —4 —3 —5
—5 —6 —5 —4 —4 —4 —4 —6 —7 —8 —7 —-8 —6 —4 —2 +0
+2 +4 +5 +6 +7 +8 +9 +10 +10 +11 +11 +12 412 +11
1D --10\--8 --5 --2 —2 —3 —7.
gu M: to +1 +2 +2 +2 +2 +2 +3 +4 +2 +1 —1I —2 —3 —3 —3
Sea i ee Me eer tape ee? em Wi ime ec ta once ure) Fa
+o +o +o —1 —2 —3 —3 —4 —4 —4 —4 —4 —4 —4 —4 —4
aoe EOE 4-2 2) 3 Ao EO oO ay ey eg
+7 +6 +5 +4 +3 +2 +2 +1 +1 +1 +0 +0 —1 —2 —2 —3
—4 —4 —4 —4 —3 —2 —I —1 +o +o =o.
The 12-month period of terrestrial causation
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.

+01 +06 —2.1 —6.7 —0.0 +1.7 +1.4 +2.1 +4.3 +6.2 +13.2 +13.5

16 _ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

dental errors of observation will still load the curve with their saw-
toothlike vibrations about its true course. No system of periodicities,
which may truly represent the true courses of the solar variation, can
possibly follow these small accidental errors of observation. It is
therefore unreasonable to demand that such a system of periodicities,
even though the true one, can be expected to reduce the average devia-
tion of its curve from the curve of observation below the one-tenth

TABLE 3.—Average departures of synthetic from observational curve

Aug. 1920—Mar. 1922, 20 months, 2.01 tenths percent.
Aug. 1923—July 1926, 36 months, 1.82 “ >

Aug. 1926—Dec. 10945, 233 months, 1.10
Jan. 1945—Dec. 1950, 60 months, 2.38
Aug. 1920—Dec. 1950, 349 months, 1.45

“cc 6c

of a percent found. For though, as stated, the probable error of first-
rate 10-day means, as found by comparing the simultaneous observa-
tions of two solar-constant observations, is 1/25 percent, very many
10-day means are not first rate, as explained above. Moreover the
“average deviation” is 5/4 of the “probable error,” as is well known,
raising the figure to 1/19 percent for the average deviation of first-
rate 10-day means.

The real crux of the question, as between the hypothesis of constant
solar radiation, and solar radiation varying in 23 regular periods,
painstakingly determined and tested by several criteria of reality, lies
in considering the large excursions of the curve of observation from
its mean. Examples of such methodically marching excursions are
found from 1924 to 1927, from 1929 to 1933, from 1937 to 1942, and
from 1947 to 1949. The hypothesis of a constant solar radiation offers
no explanation for them. On the other hand, the synthetic curve fol-
lows these large, methodically marching excursions with some fidelity.

Yet notwithstanding this striking harmony in the principal features
between the curve of observation and the synthetic curve of regular
periodicities, there are limited intervals of substantial disagreement.
Among these the major one occurs in 1922 and 1923, regarding which
I have already written. The disagreement in 1920 and 1921 may be
attributed to the incomplete development of the short method of solar-
constant determination in those earliest years. The same perhaps
applies to the disagreement in the years 1924 and 1925, for even then
the short method was not fully developed, as now used. As for the
period 1946 to 1950, Mr. Aldrich inclines to think the scales of
pyrheliometry may have varied a little in those years. There is also

NO. IO SOLAR-CONSTANT PERIODICITIES—ABBOT 17

a possibility that, in carrying the computations so far forward as 1950
from their base in 1920, slight errors in the length of the periods have
accumulated so as to mar the results of synthesis.

Brief intervals of unusually large divergence between the syn-
thetic and the observed curves occur in 1927, 1929, 1934-1935, 1938,
1940-1941, and 1944. Nearly all these cases occur at the times of
the year when sky conditions for observing are inferior at one or both
stations, as indicated by figures 7 and 8, pages 70 and 71, Annals,
volume 5, already cited. It is not probable, however, that regular
periods of variation include all the variations of solar radiation. We
know, indeed, that outbursts of sunspots and flares cause changes in
the sun’s output of radiation. Some of the discrepancies referred to
are doubtless due to such causes.

I hope the reader will agree that the synthesis of 23 independently
and separately computed periodic terms has represented, to within the
error of observation, the march of the solar constant as given by the
monthly means of the original observations from 1920 to 1950, ex-
cluding the extraordinary values of 1922 and 1923. This close agree-
ment in form and amplitude between the observed and the synthetic
curve seems to me a fourth kind of evidence supporting the existence
of a complex of over 20 regular periods all approximately integral sub-
multiples of 272 months in the observed variation of the sun’s output
of radiation.

It will occur to the reader that curves of solar observation should
tend to repeat their features after 272 months, or approximately 23
years. There is a slight indication that the curve of 1921 in figure 4
is similar to that of 1944, but the work of 1921, as mentioned else-
where, is too inaccurate to prove it. In the years 1922 and 1923 oc-
curred a unique large depression of the curve of observation. A real
test must begin with the year 1924. Unfortunately, as stated else-
where, there appears to have been a change of scale of about 4 percent
in 1948. To correct for it, I subtract 32 units from all the monthly
means, July 1948 to February 1950.

In figure 4A, I superpose the corrected curve 1947 to 1950 (light
line) upon the observed curve of observation 1924 to 1927 (heavy
line). The similarity is striking. During 48 months there are five
large divergencies: 0.55, 0.50, and three of 0.45 percent. The ex-
treme range of the great feature shown in figure 4A is 0.9 percent,
and the average deviation between the curves is but 0.19 percent—
less than the expected combined probable errors of observing. One
regrets that the interval, 276 months, exceeds the expected interval,

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

272 months. But as solar conditions modify the lengths of the sun-

spot cycles, they may also slightly modify that of the 272-month cycle
from time to time.

\
Hote

Je ih
; ty

Fic. 44.—Comparison of solar weed 1924-1927 phe lines) and
1947-1950 (light lines).

SCALE OF SOLAR CONSTANT NEARLY UNCHANGED IN 30 YEARS

It is very pleasing that the comparison of synthesized and original
curves shows the features generally with equal amplitudes in the two
curves. The comparison gives no indication that the scale of observa-
tion has changed in 30 years, except perhaps for a rise of 3/10 percent
from June 1948 to January 1950. This is remarkable in view of many

changes of instruments and of procedures that have taken place mean-
while.

NO. IO SOLAR-CONSTANT PERIODICITIES—-ABBOT I9

APPENDIX 1
SOLAR-CONSTANT MONTHLY AND 10-DAY MEANS, 1920-1950

Doubtless there are those who are engaged in research on cycles in
various lines who may wish to know the Smithsonian results on solar
variability as nearly as possible up to date. Mr. Aldrich kindly permits
me to publish the following table (table 4) giving the percentage ex-
cesses of solar-constant values above 1.900 calories from 1920 to 1950.
These percentage excesses are in the form of means of Io days (1.e.,
decades of months) and means of months. Taking the first trio of
values, given here for illustration, the table may be explained as
follows. We have:

Zoe eco. Tea mat
2) Ge WEL Os. 258839). 253°
2: (Oy EEE O, 3, 205

The above figure 2, with the figure 0, makes 20, meaning the year 1920.
The figure 8 means August, the eighth month of 1920. The Roman
numerals I, II, III stand for the first, second, and third decades of
August. That is: August I-9, 10-19, 20-31. The values 154, 139,
165 represent decade-means of the daily excesses of the solar constant
by which these observations exceeded in ten-thousandth parts of the
mean solar constant (taken as 1.94 calories) the value 1.9000 calories.
Thus the value 154 signifies that the mean solar constant for the first
decade of August 1920 was 1.54 percent of 1.94 or 0.0299 calorie
above 1.90 calories. Finally, the value 153 is the mean of the three
decade values and signifies that the average solar constant for August
1920 was 1.90+1.53 percent of 1.94 calories, or 1.930 calories.* As
stated above, the percentages of excess over 1.90 calories was chosen
to suit my investigation because, first, all values are positive, and
second, results come out in percentages of the solar constant.

APPENDIX 2
PROBABLE SOLAR-CONSTANT VALUES BEFORE 1920

Smithsonian solar-constant observations were made in the summers
on Mount Wilson, Calif., in most years from I905 to 1920. But
partly because of experimental crudity, and partly from the variability
of sky transparency, and mainly because those measurements were
all made by the fundamental “long method,” which requires constant
sky transparency for hours, the results were wide-ranging, from about

8 This result is far out of line, and indicates experimental error. In drawing
figure 4 I have assumed, instead, 235, given in parenthesis in table 4.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

WOON ANARW HH

154

139(235)

165
263
227
227
227
278
206
278
258
201
2904
263
278
299

TABLE 4.—Ten-day and

153

239

237

246

278

294

268

249

242

259

201

259

223

253

290

205

263

220

235

192

156

108

100

122

Ais

lee:

monthly means

273
242

283

129

118

190

126

165

158

182

ISI

184

211

249

220

VOL. 117

278
283
288
299
263
258
263
252
216
221
258
247
237
247
258
263
278
206
258
221
273
263
263
242
232
237
232
216
252
242
247
237
258
237
258
196
201
206
180
211
232
Ig!
170
201
216
201
206
211
196
216
211
221
211
211
232
232
252
216
216
227
201
206
180
185
185
201
170
201
IQl
206
196
IQ!
175
242
160
154
216
216
247
232
201

283
273
244
242
247
249
251
256
234
237
247
230
196
Zane
196
206
208
214
239
220
196
190
187
198
192
195

227

NO. IO SOLAR-CONSTANT PERIODICITIES—ABBOT 21

TABLE 4.—Continued

Zoe I,7 244 206 2, Sepulioy G25) 20r 3.0L I,1 406 221
II 245 242 It 326 206 II 407 232

III 246 216 221 III 327 206 204 iil 408 237 230
2. 621,97. 247) 258 25, One als Oo) S25e2nr 3, 12a) lym) 400) 228
II 248 221 iE 329 IQ1 II 410 247

Til 249 227 235 Til 330 211 204 Til 4II 242 237
2, 7 I, 7 250° 232 2,10 Nyon Sarr2rr cas ya 472) 242
Lith 251 216 II 332 216 II 413 242

III 252 232 227 III 333 175 201 Til 414 221 235
2) Sena le 7 SSav 211 25 LE I,9 334 206 3542 I, 2 435 227
II 254 221 II 335 227 II 416 232

IIl 255 232 227 Til 3360 237 223 III 417 165 208
2,°9) 1,7 256) 237 2,12 I,9 337 237 3; 3) welve, angers
II 257 258 II 338 237 II 419 221

IIT 258 247 247 III 339 227 234 III 420 206 201
2,10 1,7, a59 221 35, Tha lanO, 3408 200 2. Aaselazs AZIVIOE
II 260 206 TEL 341 232 isl 422 221

Iil 261 211 213 III 342 232 225 III 423 211 208
2, U0 [a7 262) 232 aun eee lou S4qeauE 25 85 Tia, 4249232
II 263 232 II 344 232 II 425 227

III 264 247 237 TANT 345 247 230 III 426 154 204
202 T7 265) 242 NE} T,0 346 232 3, 6 I,2 427 206
1A 266 227 II 347 211 Il 428 221

Ill 267 201 223 IIT 348 216 220 Til 429 221 216
at 1,8 268 221 30 Aw oly Oy 2407227 3). Felodec. Aan) 203
1 269 106 II 350 206 II 431 206

TIT 270 216 21% III 351 227 218 Tit 432 216 228
Za applarce 27 Te 237 3; Sy pelo) S52) 232 3, 8 I,2 433 106
II 272 211 II 353 252 II 434 227

III 273 201 216 III 354 242 242 III 435 216 213
25 134) Loh 2749237 A Obese lnOn 46ce242 3; Guaclazn asouror
II 275 247 II 356 273 II 437 232

Ill 276 221 235 III 357 258 258 III 438 237 220
2, 4 I,8 277 216 Say T,o 358 232 3, 10 I,2 439 211
II 278 227 II 359 278 II 440 180

III 279 242 228 IIT 360 273 261 Tit 44I 201 197
2, 5 1,8 280 227 3, 8 ).l,0) 361 242 3) Tle wig2s 4dsears
IT 281 263 iE 362 268 II 443 185

III 282 247 246 III 363 252 254 Til 444 201 199
2, 6 I,8. 283 247 3, 9 1,0 364 247 3, 12), 12) 4ase25s
IT 284 278 II 365 227 II 446 237

III 285 232 252 III 365 232 235 Tit 447 211 235
2, 7 1,8 286 232 3,10 I,o0 367 227 3) Teolise 44sn255
II 287 221 TI 368 237 II 449 247

III 288 216 223 IIT 369 247 237 Ill 450 268 258
2, 8 1,8 280 ror 3,11 I,0 370 242 3, 2a elasy 45reccs
IT 2900 227 II 371 242 10 452 242

TIT 291 227 215 III 372 263 249 Tit 453 242 247
2, 9 1,8 292 201 3,12 I,o 373 268 By Sapo nce asaeesy,
IT 293 237 if 374 278 II 455 206

III 204 106 211 i 375 263 270 III 456 206 216
2,10 1,8 295 227 3, 1 I,1 376 216 3) 4a we lae aeye2nn
II 2096 232 II 377 247 IT 458 227

IIT 207 211 223 III 378 268 244 AMER 459 I9gI 210
2ort, Jit. 8) 208.227 3. 2elolutn 9708247 3, 5a ols3. 4605106
II 299 252 II 380 258 TT 461 206

IIT 300 237 230 IIT 381 216 240 Tit 462 232 211
2,12 1,8 301 237 3) aula S2h227, 3; 6) 2h, 2) 4634206
II 302 227 co 383 237 IT 464 216

III 303 252 239 IIT 384 258 241 TIT 465 232 218
2; Tee Lo, 304,242 Kae ice Mae EE Bey 3, 7. .\.3 4664247
IT 305 258 II . 386 237 eli 467 242

AG 306 237 246 TIT 387 237 237 ITT 468 221 237
2, 2 I,9 307 232 By) hanna dh Sace255 3, Sapulyse 4604227
II 308 211 IT 389 258 II 470 221

TIt 3090 196 213 IIT 390 258 268 TIT 471 216 21090
2:13. ..1,6 310.242 3, 6 I,1 39% 247 35 Ompalaa Ay2aas2
ue 3II Ior IT 392 247 II 473 247

III 312 206 213 IIT 303 232 242 TIT 474 263 254
2) 7-4, JQ. 313) r0x 35 Wenelath 3044237 3,10 1,3 475 263
IT 314 242 TI 305 252 IT 476 237

III 315 227 220 III 306 247 245 IIT 477 263 253
2 Sw wl Ob arGe216 2) eCmpelate B070247 3,11 I, 3, 478 242
IT 317 216 II 398 258 TT 479 263

III 318 227 220 IIt 309 232 246 TIT. 480 273 250
20h ml, Ou are 200 3, \On sala te 4oo12s2 3,12 1,3 481 268
II 320 175 IT 401 232 TI 482 252

IIT 321 196 192 TIT 402 263 249 IIT 483 258 2590
2. ib ee aOe (3220206 2. TOvae lady ont 2os ae T,4 484 258
TT 323 211 II 404 221 iit 485 237

III 324 216 211 III 405 237 240 IIt 486 247 247

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS

3, 2° 1,4 487 263
II 488 216
Iil 489 227
3, 3 1,4 490 247
II 491 221
Iil 492 258

II 494 221
Ill 495 221
3, 5 1,4 406 227
II 497 242
III 498 221

II 500 258
III 5OI 252

II 503 232
Ill 504 232
3, 8 I,4 505 211
II 506 237
III 507 227
3, 9 1,4 508 232
II 509 247
Til 510 263

II 512 268
Ill 513 263

II 515 263
III 516 258

II 518 258

III 519 247

3 or 1s) S20 242
II 521 268

III 522 232

a2 seal Se s2queas
II 524 237

III 525 216

Spa eely si S20u220
II 527 242

Tit 528 263

Bra ol Se S20 as7
II 530 242

III 531 227
Sess seared 7
TI 533 232

III 534 247

ay WO AE Gu Ey Sey,
II 536 237

III 537 247

Sy ye He, Ses erg
II 539 227

Til 540 232
Beets Saruad7
II 542 263

III 543 237

3, 9) Ls) S44ii2g2
II 545 237

III 546 227

3; 10) 10,5) 547. 237
II 548 242

III 549 242

3,10 Poets) 550 247,
TI 551 268

IIT §52 252

8, T2115 553247
II 554 263

III 555 273

II 557 263

EET 558 227

a, 92) 1,16") 550242
TI 560 263

III 561 106

253 1,6 562 201
II 563 237

III 564 232

Heh ols Gh amex
II 566 237

TIT 567 237

235

242

225

230

251

241

225

247

265

263

258

247

230

242

235

242

240

235

249

232

240

256

261

242

234

223

235

TABLE 4.—Continued

237
232
247
258
252
247
252
242
242
232
252
242
232
252
263
252
252
242
268
273
258
278
263
247
247
273
242
247
237
252
211
221
227
201
211
216
180
227
237
237
237
242
221
227
232
242
232
242
252
247
237
242
227
242
247

3 247

247
258
252
278
232
268
206
221
237
273
268

5 237

242
232
206
237
237
227
206
211
227
232
227
232
221

239

252

245

242

249

266

263

254

245

220

209

215

239

227

239

245

237

247

263

235

244

249

225

223

227

VOL. II7

230

237

252

268

263

240

2i1I

223

242

NO. 10 SOLAR-CONSTANT PERIODICITIES—ABBOT 23

TABLE 4.—Continued

II I 730 252 2 I 811 216 a I 892 237
II 731 227 II 812 242 II 893 237
III 732 216 232 Iil 813 232 230 Ill 894 242 239
12 u 733 242 3 I 814 206 6 I 895 232
II 734 258 If 815 247 II 896 227
Til 735 258 253 Iil 816 221 225 Til 897 227 229
I I,r 736 232 4 I 817 211 7 I 898 258
IL 737 268 II 818 227 II 809 242
Ill 738 242 247 III 819 237 225 III 900 242 247
2 I 739 216 5 I 820 242 8 I QOI 247
II 740 247 II 821 232 II 902 232
III 741 273 245 III 822 237 237 III 903 221 233
3 I 742 258 6 I 823 252 9 I 904 232
II 743 232 1a 824 263 II 905 216
BE 744 268 253 Til 825 263 2590 III 906 211 220
4 I 745 247 7 I 826 252 Io I 907 232
II 746 242 II 827 221 II 908 221
III 747 221 237 III 828 257 243 Iil 909 206 220
5 I 748 247 & I 829 247 II I gio 258
II 749 216 II 830 242 II QII 242
III 750 263 242 III 831 257 249 III QI2 252 251
6 I 751 263 9 J 832 232 12 I Q13 221
II 752 216 II 833 247 II 914 216
III 753 268 249 Ill 834 242 240 IIT 915 216 218
7 I 754 258 10 I 835 232 4, 1X EG 916) 22%
II 755 268 II 836 247 II Q17 227
III 756 283 270 TII 837 232 237 III 918 258 235
8 li 757 283 II I 838 237 2 I 919 211
II 758 252 II 839 247 II 920 201
III 759 242 250 III 840 206 230 III 921 201 204
9 I 760 278 12 af 841 232 3 I 922 191
II 761 263 II 842 227 II 923 185
III 762 216 252 III 843 263 241 III 924 206 194
TO! ya 763 273 4,0 1, a 844s 252 4 I 925 252
II 764 258 II 845 242 II 926 242
1090 765 247 259 III 846 227 240 Til 927 ZIT 235
Tee 766 268 2 wil 847 257 5 0 928 252
II 767 247 II 848 263 II 929 252
Til 768 247 254 III 849 252 257 Ill 930 227 244
TZN ent 769 263 a0 WET 850 216 Gy eal 931 258
II 770 258 II 851 242 Ir 932 247
III 771 283 268 IIt 852 227 228 Til 933 258 254
rt I,2 772 288 7g 853 216 7 al 934 237
II 773 237 II 854 227 II 935 258
III 774 247 257 III 855 227 223 III 936 221 239
2 al 775 247 Gye at 856 242 458) pol Om Oa7 zor
IT 776 252 II 857 237 II 938 216
Til 777 247 249 III 858 227 235 seat 939 252 230
Shas 778 221 6a 859 237 Oy cur 940 252
II 779 221 II 860 221 II 941 232
IIT 780 216 219 III 861 227 228 It 942 221 235
Zee | 781 227 7 I 862 237 ROME 943 216
IT 782 237 II 863 227 IT 944 232
III 783 232 232 III 864 237 234 oe 945 237 228
Bhi peal 784 227 4, 8 1,4 865 263 Mh veges 946 247
TT 785 242 II 866 216 ur |g 258
IIT 786 257 242 III 867 206 228 i 948 263 256
6) 'T 787 257 go. eat 868 206 a It 949 304(?)
It 788 237 II 869 221 we 3 BzER ee
TIT 789 247 247 III 870 191 206 t gsr 221(?)
Pd Sox 790 252 may al 871 232 aaa 1?’ 952 278
II 791 257 II 872 206 953 258
III 792 232 247 III 873 185 208 III 954 258 ahs
Bat, 2 703 242 II I 874 247 2 I 955 216(?)
Il 704 237 II 875 237 TE = 956 237
TIT 795 237 235 III 876 232 239 IIT 057 242 232
Bis eal 796 247 TAL 877 237 Seal 958 185
II 797 106 II 878 268 II 959 206
TIT 798 232 225 IIt 879 232 246 Til 960 216 202
ios eal 799 232 4 i <,5 880 2or 40 ak 961 191%
TI 800 227 13) 881 arr II 962 227
TIT = Bor 216 225 III 882 206 206 III 0963 232 217
ik 802 237 20m 883 242 Seed 964 232
TI 803 242 TI 884 247 IT 965 232
III 804 257 245 IIT 885 232 240 Til 066 216 227
r2) nT 805 232 3 I 886 232 Gyan 0967 211
II 806 211 TI 887 232 II 968 247
III 807 227 223 It 888 206 223 IIt 969 242 233
I T,3 808 206 4 I 889 232 7 I 970 237
TI 809 206 II 890 253 TI O71 232

Tit 810 211 208 TIT 891 247 244 Ill 972 211 227

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLs LL7

TABLE 4.—Concluded

8 I 973 216 10 I 1015 258 12 I 1057 263
II 979 221 II 1016 258 II 1058 242
III 975 252 230 Iil 1017 253 356 IIl 1059 288 264
9 I 076 221 II I 1018 278 Bee be I,o 1060 288
II 977 237 II 1019 268 II 1061 268
IIL 978 221 226 Til 1020 283 276 Iil 1062 268 275
Io i 079 227 12 I 1021 278 2 I 1063 227
II 980 237 II 1022 283 II 1064 237
Tit 981 263 242 Tit 1023 278 280 Til 1065 247 237
rye I 982 273 As ot I, 9 1024 278 3 I 1066 227
II 983 242 II 1025 247 ih 067) 227
III 984 247 254 III 1026 273 266 III 1068 211 222
12 I 985 258 2 I 1027 304 a I 1069 221
II 986 237 II 1028 2094 II 1070 211
III 987 263 253 III 1029 232 278 III 1071 206 213
45,00 I,8 988 278 3 I 1030 242 5 I 1072 232
II 989 247 II I03I 252 II 1073 206
III 990 263 263 Iii 1032 206 233 III 1074 247 228
2) Wak 991 273 4 1 1033 221 6 SLE S075 242
Wi 992 268 II 1034 206 II 1076 232
III 993 258 266 III 1035 258 228 III 1077 247 240
3. pt 904 278 5s I. 1036 242 7 I 1078 211
II 9095 247 II 1037 273 II 1079 242
III 996 247 257 III. 1038 242 252 III 1080 232 228
45 pee 097 203(?) 6 I 1039 232 5, 8 1,0 1081 253
II 998 252 II 1040 ror II 1082 253
III 999 258 268 III IO4I 242 222 III 1083 216 241
5 tly Ser0008258 7) Wale OA 2 287 o @6©1-~—s1084 206
II 1001 252 II 1043 242 Il 1085 237
III 1002 263 258 TII 1044 247 242 III 1086 237 227
6 I 1003 283 8 I 1045 242 10 I 1087 243
II 1004 273 II 1046 263 ie 1088 249
IE | roo5 273) 276 III 1047 221 242 III 1089 283 258
7 I 1006 299 9 I 1048 227 II I 1090 263
II 1007 288 II 1049 237 II 10QI 227
TIT 1008 283 290 III 1050 206 223 III 1092 252 247
4, 8 1,8 1009 283 10 IT ros1 232 12 i, “woos, 232
II 1010 278 II 1052 221 II 1094 247
III ror 237 266 III 1053 263 2390 III. 1095 227 235
9 I IOI2 252 II I 1054 268
II 1013 278 TT 1055 247
III 1014 278 269 TII 1056 237 251

1.9 to 2.0 calories, or even more. Still, by forming these less-accurate
solar-constant values into large groups of days, according to magni-
tude, H. H. Clayton was able to correlate solar changes with weather
elements.®

It now occurs to me that since the periodicities now discovered in
the solar emission have been expressed as to form and amplitude,
and since 1920 seem to be permanent as far as known in period, ampli-
tude, and form, it may be worth while to synthesize monthly mean
solar variation backward from 1920. This done, it would be possible
to compare the values synthesized with monthly mean solar-constant
values observed on Mount Wilson. If, on the whole, high, medium,
and low solar constants as synthesized correspond to high, medium,
and low Mount Wilson values, it will be a confirmatory evidence of
the sun’s real variability, of the constancy of periodicities, of their
comprising nearly the total solar variation, and of the value of Clay-
ton’s work on the correlation of solar variation with weather.

Table 5 gives the synthesized monthly solar-constant values from

9 Smithsonian Misc. Coll., vol. 68, No. 3, 1917.

NO. IO SOLAR-CONSTANT PERIODICITIES—-ABBOT 25

August 1908 to December 1920. These results are given graphically
in figure 5,C. These are actual estimated solar constants in calories
per square centimeter per minute, not, as in table 4, percentage de-
partures from 1.90 calories.

COMPARISON OF SYNTHETIC WITH MOUNT WILSON SOLAR-
CONSTANT VALUES

From table 53, page 193, volume 4, Annals of the Astrophysical Ob-
servatory, I take monthly solar-constant values determined from
Mount Wilson observations in the months May to November, 1908
to 1920. I omit four values, July and August 1912, because the sky
was then very much fouled by dust from the volcano, Mount Katmai.’
I also omit July values of 1910 and 1917 because they are very wild
indeed, far beyond the limits of dispersal of the others.

Having plotted the Mount Wilson values and such parts of the
synthetic series as corresponded in time with them, I saw that there
was a gradual rise in values in both observed and synthetic series from
1908 to 1914. I drew straight lines best following this trend to repre-
sent the means of the values over that interval, and read off the de-
partures of the individual solar-constant values on the plot from
these lines. For the rest of the total interval, that is 1915 to 1920, I
read departures from straight horizontal lines drawn in the mean of
ordinates. The plot was in arbitrary units, with the units for ordinates
in the synthetic plot twice as large as those for the Mount Wilson
data. These departure values follow in table 6.

Taking the sums of the data in the columns of table 6 they yield:

Mount Wilson +synthetic= a =1.77. Recalling the ratio of units,
2 to I, it appears that the dispersal of Mount Wilson data is 3.54
times as great as that of the synthetic data. The synthetic curve 1920-
1950, however, as plotted in figure 4, shows practically the same range
of variation as does the curve of original modern observations. Hence
it appears that the Mount Wilson solar-constant observations of 1908
to 1920 are probably 34 times less accurate than the modern work set
forth in table 4.

Taking account of the numbers of departures of the same sign in
the columns of table 6, and the numbers of them of opposite signs,
the sums are 28 and 21.

Taking the sums of departures that are of the same sign in both
columns, the results are 324 for Mount Wilson and 170 for the syn-

10 See Smithsonian Misc. Coll., vol. 60, No. 20, 1913.

1908 Aug.
Sept.

Oct.

Nov.

Dec.

1909 Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
1910 Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
Io1t Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.

SMITHSONIAN

TABLE 5.—Synthesized solar constant, 1908-1920

Values to be prefixed by 1.9

IQI2

1913

1914

Jan.

Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.

1915 Jan. 45

1916

1917

Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.

43
46
50
48

MISCELLANEOUS COLLECTIONS

VOL, 117

1918 Jan. 47

1919

1920

Feb.

Mar.

Apr.
May
June
July
Aug.

Sept.

Oct.

Nov.

Dec.
Jan.
Feb.

Mar.

Apr.
May
June
July

Aug.
Sept.

Oct.

Nov.

Dec.
Jan.
Feb.

46
43

SOLAR-CONSTANT PERIODICITIES—-ABBOT

NO. 10

* (Sa}eUuIpsO ) oneyiuds "SA (QBSSIOSqe) UOS|IAA JUNO ‘UksU 34} WoIJ soinjiedep Jue}suOD
-Jejos Ajy}JUOW UveU Jo UOsIIedWIOD YW ‘q ‘OZ6I 0} QObI ‘sat}IOIpotsiad Ez wWo1, pozisayyUAs se ‘satIoyed UL sonTeA yUR}sUOD-Ie[OSG ‘Q—S ‘oly

PiRB@EREERRBERS)
AA VANS
HW A ee

OvG'!

CUE L Le
RSARBEAZERAESTE SC
SZBSeeS2ask2ets=
SERRE SESese

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7
thetic data. The corresponding sums for departures of opposite signs
are 199 and 135. Thus, according to Mount Wilson, agreeing de-

partures preponderate in total magnitude over disagreeing depar-

TABLE 6.—Comparison of Mount Wilson and synthetic values

Mount Mount Mount
Wilson Syn. Wilson Syn. Wilson Syn.

1908 IQ12 I916

Aug. +45 +16 May +5 +1 June “— 7 — 6

Sept. +28 +15 June — 8 — I July —3 —I10

Oct. +43 +13 1913 Aug. + 2 —18
1909 Aug. —7 +65 Sept. — 8 —I2

June +17 — 6 Sept. —30 + 3 1917

July —3 —I I9I4 July +20 — 6

Aug -+12 +1 June +0 —7 Aug. -+ 6 —2

Sept. — 7 —I July +4 —I5 Sept. — 2 —4
I9IO Aug. -+II —I4 1918

May +12 + 8 Sept. —II —I4 June —7 + 2

June — 5 +7 Oct. — 6 —I5 July +4 =) (0)

July 24, E45 1915 BAM! t= 3) one

Aug. —II +7 June — 8 +o Sept. + 9 + 4

Sept. —13 +65 July — 3 — 6 1919

Oct. + 3 +65 Sept. + 1 —I4 June +7 +4
IQII Oct. +17 —I10 July $= so —2

June +15 + 8 Aug. — 5 —2

July —13 +5 Sept. — 6 + 6

Aug. —2 + 3 1920

Sept. + 6 +1 July —25 — 6

Oct. —17 —2 Aug. — 3 — 8

Sept. —37 — 6

tures as SA =16, Similarly, for synthetic values the results are
nad =1.3.
135

Finally, I show in figure 5,D, the Mount Wilson departures as
abscissae against the synthetic departures as ordinates. The plotted
points are greatly scattered, as the inaccuracy of Mount Wilson
solar-constant values would lead us to expect. Yet, on the whole, the
comparison indicates that high departures tend to occur simultane-
ously in both sets of data, and low departures similarly.

Thus four kinds of rough indications agree to confirm the view that
the synthetic solar-constant values of 1908 to 1920 are supported as
to their validity, at least in some degree, by the evidences from Mount
Wilson observations. The four evidences are: 1. Both sets of data
yield upward trends from 1908 to 1914. 2. Departures from repre-
sentative lines have the same signs 28 times, opposite signs, 21.

NO. I0 SOLAR-CONSTANT PERIODICITIES—ABBOT 29

3. The summation of departures of the same sign exceeds that for
those of opposite sign about 14 times. 4. The plot of departures in-
dicates a positive correlation between Mount Wilson and synthetic
solar-constant values.

The great inferiority in accuracy of Mount Wilson values of the
solar constant forbids a high degree of correlation, even if the syn-
thetic values are as correct from 1908 to 1920 as they are from 1920
to 1950. This inferiority arises from the fact that all the Mount
Wilson values result from observations by the “long method.” That
method requires for accuracy a sky of constant transparency over
several hours. If the sky improves, the solar-constant value is too
high, and vice versa. Moreover, only one value was obtained per
day with the “long method.” In modern solar-constant work by the
“short method,” several values are obtained and combined on each
day of observation. The sky is required to retain uniform trans-
parency only during about 10 minutes of each observation. It might
vary decidedly from one determination to another of the day’s group,
and yet all the solar-constant values of the day be closely agreeing.

SOLAR CONSTANT AND SOLAR CONTRAST

The Mount Wilson work offers another test of the probable validity
of the synthetic solar-constant curve of 1908 to 1920. From 1913 to
1920 we were accustomed to produce drift energy curves in several
wavelengths, observing intensities along the east-west diameter of an
8-inch solar image, on every day that we observed the solar constant
of radiation. These U-shaped curves, which show the contrast in
brightness between the center and edges of the sun’s disk, were all
measured as described in volume 4 of the Annals of the Smithsonian
Astrophysical Observatory. We used an empirical formula to obtain
a value to represent the average contrast between center and edge of
the sun’s disk on each day of observation. These data are given in
tables 75 to 82 of volume 4 of the Annals.

It was thought probable that the “solar contrast” would be greater
on days when the “solar constant” was higher. Some figures, indicat-
ing that this is so, are given in volumes 3 and 4 of the Annals.

Table 7, which follows here, is prepared from the “‘solar contrast”
tables of the Annals, volume 4, and from table 6, just given, which
presents synthetic solar-constant values of 1908 to 1920. To prepare
the solar-contrast values for this use, means of the daily values are
taken of every month given in Annals 4. Then, in order to eliminate
systematic errors which might introduce inconsistencies, a separate

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL SEL

mean value is computed for the available months of each year, 1913 to
1920. Differences from these yearly means are given in column 2 of
table 7. To make the synthetic solar-constant values entirely com-

TABLE 7.—Comparison of synthetic solar-constant departures with solar-contrast
values of 1913-1920

Solar-constant departures in thousandths of a calorie.

Solar Solar
constant contrast
+17 +19
= 3 —32
Se +14
+36 —35
+ 6 —24
—34 —18
—I4 +28
70 +49
+20 +10

ta) —29
—40 —— 5
—I10 (s)
+30 ao
—I7 —18
aia 4-15
=aNiy +14
—I4 —23
—4 —I12
+ 6 + 8
+16 +16
+5 —8
—I5 —I3
S15 eed
+25 +40
ee +46
aan +18
as Sahu
+23 13
al sale:

parable to these contrast values, separate means of them are taken for
each year of the comparison, including only the months used in ob-
taining the separate contrast means. Differences from these synthetic
solar-constant means, expressed in thousandths of a calorie, form
column 1 of table 7.

Counting the numbers of months when values in columns 1 and 2
have the same sign and opposite signs, the numbers (counting zero

NO. 10 SOLAR-CONSTANT PERIODICITIES—ABBOT 31

values into each group) are 18 and 13, respectively. So here is another
straw pointing to the reliability of the synthetic solar-constant values.
But more convincing, and more informing, is figure 6. Here the

-20

Fic. 6.—Mount Wilson solar contrast (abscissae) vs. synthetic solar constants
(ordinates).

values in the columns of table 7 are plotted against each other, solar
constants as ordinates, solar contrasts as abscissae. In order to bring
out plainly the fact that higher contrast values attend higher synthetic
solar-constant values, stars 1, 2, 3, 4, 5, have been plotted to give the
centers of gravity of groups of 8, 8, 5, 5, and 2 months, respectively.
A full heavy line has been drawn to show the trend of the results.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 11

Roebling Fund

IMPORTANT INTERFERENCES WITH NORMALS
IN WEATHER RECORDS, ASSOCIATED WITH
SUNSPOT FREQUENCY

BY
C. G. ABBOT

Research Associate, Smithsonian Institution

(Pustication 4090)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
MAY 20, 1952

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Roebling Fund

IMPORTANT INTERFERENCES WITH NORMALS
IN WEATHER RECORDS, ASSOCIATER
WITH SUNSPOT FREQUENCY

By C. G. ABBOT
Research Associate, Smithsonian Institution

World Weather Records + comprise long tables of monthly weather
records of pressure, temperature, and precipitation. At the boitom of
each table are given normals of these elements, determined by averag-
ing the long columns of monthly values for each month of the year.
By subtracting these normal values from the numerous observed
temperature values, tables of temperature departure are obtained. By
dividing the numerous observed precipitation values by appropriate
normal values, percentages of normal precipitation are tabulated.

By using percentages of normal precipitation thus obtained in a
study of the effects of periodic solar variations on the precipitation of
Peoria, Ill., important influences of sunspot frequency were un-
covered. In the first place, the average percentage precipitation was
about 9 percent higher when Wolf sunspot numbers exceeded 20 than
when the Wolf numbers were below 20. In the second place, the
average computed percentages showed large 12-month periodicities.
The amplitude of these periodicities is about 8 percent when Wolf
sunspot numbers exceed 20, and about 16 percent when the Wolf
numbers are below 20.

The 12-month periodicities, just described, are roughly opposite in
trends. With sunspot numbers above 20, as computed, high average
percentage precipitation occurs in the first half of the year, low per-
centage precipitation in the second half. The maximum is in June,
the minimum in October. With sunspot numbers below 20, low per-
centage precipitation occurs in the first half of the year, high per-
centage precipitation in the second half. The maximum is in August,
the minimum in March.

1 Smithsonian Misc. Coll., vol. 79, 1927; vol. 90, 1934; vol. 105, 1047.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 11

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7
To determine these facts I used the normal values of precipitation
published in World Weather Records for most of the time, but for

recent years, for which World Weather Records are not available, I

ite)

100

JAN. MAR. MAY JULY SEPT. NOV.

Fic. 1.—Twelve-month periodicities in precipitation at Peoria, IIL, depending on
sunspot frequency, but hidden by using published normals.

used the normals published by the U. S. Weather Bureau, which differ
slightly. These normal values for Peoria are as follows, in inches of
precipitation.

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.

Earlier... 1:75 1:70 2.80 3:28 3:04 3.71 3:75 99:44. > 4:24 e244) $2.92) 71366
Water). 2:3y a7 9Oheit. OS 2576) Se55 e700 306mg -7 0 3°06) 4) 4.700) 2.02 uaeotmeescod

The percentages of normal precipitation were computed for me by
assistants, and I did not know until lately that two slightly different

NO. II WEATHER AND SUNSPOT FREQUENCY—ABBOT 2

sets of normals were used. In further studies I intend to compute
two sets of normals for Peoria from 1856 to 1950. One set will be
suited to sunspot numbers above 20, the other to sunspot numbers
below 20.

Separating the 82 years of the Peoria tabular monthly mean values
of precipitation, 1858 to 1939, into two groups, 50 years of high sun-
spots, 30 years of low sunspots, and 2 years being omitted as mixed,
I computed the following monthly mean percentage precipitation
values, based on the above quoted normals.’

Jan. Feb. Mar. Apr. May June
Wolf number > 20... 105.6 107.0 107.9 108.5 109.1 108.9
Wolf number < 20... 91.1 89.8 89.2 91.8 93.0 Q1.7
July Aug. Sept. Oct. Nov. Dec.
Wolf number > 20... 105.6 102.4 IOI.9 101.2 103.4 102.3
Wolf number < 20... 98.0 105.0 104.8 102.8 101.9 08.7

These results are plotted in figure 1. The curve A corresponds to
Wolf numbers > 20, the curve B to Wolf numbers <20. One readily
sees that high Wolf numbers attend higher average precipitation at
Peoria; that strongly marked 12-month periodicities are hidden by
tables of percentage precipitation as ordinarily computed from pub-
lished normals; and that the amplitudes of these periodicities are
large, and their trends roughly opposite, for sunspot numbers above
and below 20.

It is quite obvious that no sound results on periodic fluctuations of
Peoria precipitation can result from tabulating percentage precipita-
tion computed from published normals. Take, for instance, the 7-
month solar-radiation periodicity. Tabulations of numerous 7-month
columns, covering many years, would be used. One or more of these
columns would start with June in years of low sunspot activity.
Curve B of figure 1 shows that in this column there would be a
spurious increase of 13 percent from June to August, and a spurious
decrease of 6 percent from August to December. Another column
would start perhaps with August in some other year. This column
would contain a spurious decrease of 15 percent to its end in February.

If such sunspot influences are hidden in the published normals for
one weather element at one station, it is probable that similarly caused
defects are hidden in published normals for all weather elements at
all stations. Statistical meteorologists will do well to be on the watch
for them.

2 These computations employ the percentages of normals smoothed by 5-month
running means.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 12

TWO ABORIGINAL WORKS OF ART
FROM THE VERACRUZ COAST

(Wirth TuHree Pirates)

BY
PHILIP DRUCKER

Bureau of American Ethnology

e200
Cl btbbhl TPs

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C9> <s Ato

(PUBLICATION 4091)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION

AUGUST 26, 1952

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The Bord Baltimore (Press
BALTIMORE, MD., U. 8. As
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TWO ABORIGINAL WORKS OF ART FROM THE
VERACRUZ CORsa

By PHILIP DRUCKER
Bureau of American Ethnology
(WiTH 3 PLATEs)

The two specimens described here were excavated at the site of
Cerro de las Mesas, Veracruz, by the National Geographic-Smithso-
nian Institution Expedition to Mexico in 1941. Both were referred
to in the published accounts of the work.t They could not be described
adequately, however, at the time the reports were written, for both
were badly broken, and their restoration was a long, painstaking proc-
ess. This was particularly true of the pottery statue with which the
preparators at the Museo Nacional de México struggled intermittently
for several years before they found a way to bolster the fragments to
support the figure’s considerable weight. Both pieces are now among
the collections of the Museo Nacional.

The specimens are important on two counts. From the cultural
standpoint they reflect rather clearly the cultural heritage of their
makers, and as objects of art both deserve consideration. As examples
of artistic expression these two pieces rate high among the host of
finer achievements of Middle American civilizations. Therefore, the
accompanying descriptions and plates are presented for the benefit of
students of Mesoamerican archeology and also for those interested
in prehistoric American art.

THE TURTLE SHELL

The first piece to be described is a carved turtle shell (pl. 1, and fig.
1). It was found among the numerous mortuary offerings associated
with burial I1-18, in trench 30, which belongs to the Lower II period
of the site. The small shell, about 15 cm. in length, was complete with
both carapace and plastron. Six holes about the periphery of the cara-
pace suggest that it was meant to be lashed onto something, but the use

1 Stirling, M. W., Expedition unearths buried masterpieces of carved jade, Nat.
Geogr. Mag., vol. 80, No. 3, pp. 207-302, 1941; Drucker, P., Ceramic stratigraphy
at Cerro de las Mesas, Veracruz, Mexico, Bur. Amer. Ethnol. Bull. 141, pp. 9,
12, 1943.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 12

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

of the object is unknown. Perhaps it was a ceremonial rattle. Restora-
tion of the many fragments into which it had been broken revealed a
complex design consisting of a number of symbolic motifs engraved
on the carapace. The design appears to have been engraved with some

(s)
ZN

(=e
NZLLZZTLTTLL

Fic. 1—Design carved on turtle shell from Cerro de las Mesas. (From drawing
by Covarrubias. )

rather sharp implement ; then the areas adjoining the principal fields
were scraped away to shallow bevels to give an impression of bold re-
lief. The central figure of the design is the head of a man wearing an
elaborate helmet consisting of a jaguar head, surmounted by a leering
monkey skull. The jaguar’s jaws are open so that the wearer’s face
appears between them. Such headgear was often used, we know, by
Mexican priests on occasions of ritual. Considerable stylization has

NO. 12 ABORIGINAL ART FROM VERACRUZ——DRUCKER 3

taken place in the representation of the jaguar, but the beast may be
recognized by the short nose, the low-domed, catlike forehead, and the
large, round ears. The jaws have been modified until they differ some-
what from those usually seen in drawings from the area. They have
a faint suggestion of the powerful beak of some raptorial bird. The
significance of the monkey skull with its peculiar 3-tined crest is
difficult to interpret.

Framing the central figure is a band consisting of two plumed ser-
pents. The heads of these weird creatures are to be seen on either side
of the head of the central personage with the helmet. The serpents’
bodies are represented by bands covered with three parallel zones of
small elements. The outermost of these shows unmistakably the broad
belly scales, the cross hatching of the middle band representing the
smaller scales of the sides, and the upper (or inner) zone indicating
markings of the reptiles by opposed sets of sloping lines.

Good design is achieved by the contrast between the small compact
elements of the serpent bodies and the more open areas of the central
figure. There is a horizontal band set off by the plumed serpent heads,
with their sweeping curves and open spaces, across the middle of the
field that invariably leads the eye to the human face at the center, aug-
menting the emphasis of the location of the face on the high point of
the shell’s convexity. Were it not for this horizontal movement the
components of the intricate pattern would have been lost in a confused
mass of lines and small forms.

Another striking feature of the carving from the standpoint of de-
sign is its deliberate asymmetry that gives it movement and avoids the
static monotony of perfectly balanced patterns. The bodies of the
upper and the lower serpents are not in line. The left side of the lower
one is offset markedly to the viewer’s left ; the right end, where it joins
the head, is set over only slightly, producing a total effect of motion
to the left. A similar trend is produced by the monkey skull and the
plumes of the right-hand serpent which project beyond the central
band and produce, with the rest of the open curvilinear areas of the
central band, a triangle whose apex lies, or rather points, to the left—
the same direction in which the central personage is facing. The
plumes of the left-hand serpent were suppressed apparently because
their use would have arrested the consistent leftward motion of the
pattern.

A small area between the head and ear of the right-hand plumed
serpent and the lower jaw of the jaguar headdress is partially cross
hatched. It looks as though the artist had begun to fill in the entire
strip between the central figure and the frame, then stopped. Why he

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

did not complete this filler cannot be determined. It may be that he
realized such a filled area would interrupt the transverse motion of the
design.

All in all the design is well executed and obviously planned. Stylis-
tically the carvings belong to the Highland tradition. The motifs:
Personages wearing headgear representing the open fauces of an ani-
mal or monster, and plumed serpents, are common in the Highland
from Teotihuacan to Aztec times. (They occur also in Maya art but
the style is not in the least Mayan.) The treatment: A nonrealistic
stiffness of the central figure, exuberant detail and filler, remind one
strongly of paintings from Teotihuacan frescoes, and also certain
Oaxacan stelae. Neither the characteristics of detail nor the general
staid simplicity of Olmec art can be detected in the piece. Although
the design is complex it lacks the interweaving of elements, the sinu-
ous interlocked lines, of the Central Veracruz carving found on stone
yokes and palmas, the style often called (or miscalled) ‘“Totonac.”

There is a noteworthy stylistic resemblance between the carving on
the turtle shell and that of four Cerro de las Mesas stelae: those num-
bered 3, 5, 6, and 8.2 Of all the better-preserved stone monuments
from the site, these four form the most sharply defined group stylisti-
cally and, like the turtle shell, appear to derive their stimulus from
Highland influences. This similarity suggests the provisional assign-
ment of these stelae to the time horizon of the turtle shell, which, on
the basis of the pottery associated with it in the burial offerings, can
be dated as of the Lower II ceramic period.

THE STATUE OF XIUHTECUTLE!L

The clay statue shown on plates 2 and 3 was excavated by Stirling
in Trench 34. It is to be attributed to the same temporal horizon as
the turtle shell, Lower II. The large, heavy vessel that is supported on
the head of the figure had been placed on a step and was buried by an
enlargement of the mound. Portions of the body of the statue were
inside the vessel, and pieces of clay arms and legs were stood vertically
around the mouth. Within the largest fragment of the torso were
bones of a human infant. Underneath the other fragments was the
head.* In other words the object would seem to have been deliberately
broken up so that all the pieces could be piled together as an offering
during this particular enlargment of the mound.

2 Stirling, M. W., Stone monuments from southern Mexico, Bur. Amer. Ethnol.
Bull. 138, pls. 23, 24, and figs. roc, Ioa, 9, 11b, and Iic, 1943.
8 Stirling, op. cit., 1941; Drucker, op. cit., 1943, pl. 8,c,f.

NO. 12 ABORIGINAL ART FROM VERACRUZ—DRUCKER 5

The shattered condition of the specimen when found made it diffi-
cult to recognize, in the field, the being represented ; it was only when
we began trying to fit parts together that we discovered that the
bearded head actually belonged with the body fragments. Even then
it was thought that the large vessel was a pedestal for the figure. After
the figure had been restored at the Museo Nacional de México, how-
ever, Dr. Caso, Sr. Noguera, and others of the staff recognized that
the statue represents Xiuhtecutli, The Fire God (also called Huehue-
teotl, The Old Old God), and, after examining the restored vessel,
found clear indications of its former attachment to the head. After
a number of trials, a base was designed which would support the figure
adequately, along with an armature that sustains most of the weight
of the heavy vessel—really a brazier—on the head. Xiuhtecutli, in his
role as God of Fire, usually was portrayed supporting a vessel con-
taining fire in just this fashion. It should be remarked that the com-
plete figure, brazier and all, is between 80 and go cm. high, and in some
areas the clay is nearly 2 cm. in thickness. The piece is therefore ex-
tremely heavy.

The statue had not been completely restored when the photographs
shown in plate 2 were made. For that reason the flat tablets back of
the head, which helped distribute the weight of the brazier to the
shoulders, do not fit flush as they should (note daylight between the
piece and the bottom of the vessel on the viewer’s left, plate 2, b).
Likewise, the flat strips between the figure’s elbows and body and
those on the knuckles were temporary braces of wood put on till the
restoration of the figure could be completed. There should also be a
headband about the head, with a wide flat bow, painted red, across the
forehead, as in plate 3, which shows the statue as it is today in the
Museo Nacional.

The basal portion of the torso, where the buttocks should be, is
finished off square. I suspect the statue was built solidly in a bench
or step, or on an altar, to keep it from overbalancing. The upper ends
of the legs likewise are finished off square. They must have been
fastened to the bench or altar or similar architectural feature.

The restored base for the figure is made so that the left leg rests
within the curvature of the right. Actually, the left leg rested on top
of the right, the edge of the foot attaching to the upper surface of the
right leg at the point at which a small protuberance can be seen in the
photograph (pl. 2, b). This seems a most uncomfortable postition.
Perhaps it was used deliberately to increase the tension and feeling of
strain the figure portrays. On the other hand, it may have been a

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 107

customary local posture. People can get used to sitting that way, for
I have seen Marshallese do so by the hour. This is only a random ex-
ample; there is not the slightest intent to suggest any farfetched
Oceanic influences.

At first glance the figure strikes one as being extremely realistic,
but closer consideration shows it to contain a remarkable blend of real-
ism and conventionalization. The sharp-peaked indentations above and
below the eyes are completely nonrealistic, but the heavy shadows they
produce give an effect of deep sockets from which the heavy-lidded
eyes bulge. The exaggerated V-shaped cheekbones produce the effect
of sunken cheeks, making the face that of an aged person. The deeply
incised lines about the face suggest both wrinkles and lines formed by
grimaces expressing physical strain. The formless tubular arms and
legs, anatomic impossibilities, give the effect of the scrawny limbs of
an old man. The body of the figure, with its shoulders hunched and
back bent to an almost deformed angle to support a great weight, its
sagging pectoral muscles and paunch from which the elasticity of
youth have long since departed, is the most realistic unit of the piece.
The effect of great physical effort and strain is achieved through a
variety of ways, some already mentioned: The bulging eyes, the lines
of the contorted features, the angle at which the head is held and the
hunched-up shoulders, and, of course, the pronounced forward bend-
ing of the body. The original awkward position of the legs must
have completed the strained, almost tortured, aspect of the figure.

The identity of the personage as Xiuhtecutli, The Fire God, who
was also called Huehueteotl, The Old Old God, is quite clear, from
the obvious indications of age and the characteristic brazier supported
on his head. Xiuhtecutli was a Highland deity. According to Sahagun,
he was the most important of the Aztec minor gods. His cult flourished
from Teotihuacan through Aztec times in and around the Valley of
Mexico, and Vaillant found him represented in the horizon of the
Middle Cultures. His appearance at a site in the coastal plain of
Veracruz can only be interpreted as a reflection of the strong High-
land influence on the early population of the Cerro de las Mesas region.

4 Vaillant, G. C., The Aztecs of Mexico, p. 42, 1941. The small clay figurines
representing sunken-cheeked aged persons, found occasionally in the merged
Proto-Classic-Early Classic horizon of southern Veracruz and Tabasco (the
period Middle Tres Zapotes and La Venta), apparently simply represent elderly
persons, not a cult of Huehueteotl, for they lack the specific attributes pertaining
to fire, etc., that characterize that Highland deity.

NO. 12 ABORIGINAL ART FROM VERACRUZ—DRUCKER 7,

CONCLUSIONS

Both the specimens described here, with their pronounced Highland
affiliations, emphasize a point brought out by the general pattern of
ceramics from Cerro de las Mesas. Aztec influence, that is to say,
Moctezuma’s domination over the central Veracruz coast, when Cortes
and his swashbucklers landed, was but one of history’s repetitions.
Highland cultures, richly and powerfully developed in the Mixteca-
Puebla focus, had spilled down to the coast before, as demonstrated
by the abundance of Cholulteca painted pottery that marks the begin-
ning of the “Upper” culture horizon at Cerro de las Mesas. This
earlier intrusion marks a Pan-Mexican period in which cultural dom-
inance and expansion carried Mixteca-Puebla elements not only to
central Veracruz and Yucatan, but to western Mexico as far as distant
Sinaloa. But even before, during the long life of Teotihuacan culture,
geographical distance could not restrict diffusion. From the times of
the earliest occupation found at Cerro de las Mesas, the strongest in-
fluences determinable were those of Teotihuacan. Transplanted to a
new setting, some of these Mexican Highland concepts attained new
artistic heights on the Gulf coast, as the themes carved on the turtle
shell and the statue of Xiuhtecutli clearly show.

SMITHSONIAN MISCELLANEOUS COLLECTIONS Wells ail7, Wes 22, IP, at

CARVED TURTLE SHELL FROM CERRO DE LAS MESAS, AFTER RESTORATION

SMITHSONIAN MISCELLANEOUS COLLECTIONS Vols 177) Now 22; Pine

b (Bs

POTTERY STATUE OF XIUHTECUTLI, THE FIRE GOD

a, Facial detail, before restoration : b, c, statue as partially restored.

SMITHSONIAN MISCELLANEOUS COLLECTIONS Vol. 117, No. 12, Pl. 3

Cig aed

THE STATUE COMPLETELY RESTORED

Photograph courtesy National Geographic Society.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 13

“Charles DB. and Mary Waux Walcott
Research Fund .

PRIMITIVE FOSSIL GASTROPODS AND
THEIR BEARING ON GASTROPOD
CLASSIFICATION

(WitH Two P tates)

BY
J. BROOKES KNIGHT

Research Associate in Paleontology, U. S. National Museum

( PUBLICATION 4092)

| fe ath ay CITY OF WASHINGTON
sie PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 29, 1952

‘ ig
pits ui
4 4
Tenth iM
RF ”

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 13

Charles DB. and Mary Oaux Walcott
Research Fund

PRIMITIVE*FOSSIL GASTROPODS AND
THEIR BEARING ON GASTROPOD
GLASSIFICATION

(WitH Two PLatTEs)

BY
J. BROOKES KNIGHT

Research Associate in Paleontology, U. S. National Museum

(PusticaTion 4092)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 29, 1952

The Lord Baltimore Press

BALTIMORE, MD., U. & A.

CONTENTS

Page
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Rome AUC OUST CLAGIOMS «5.5 slices 5:0 Sie a.da:e o> ~ aise susan tot chwiziele tal oe saeeede eee I
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PEN Ue ATA EO ATTICSIES) alae c) ore aio: bie of otalee! ask c-oxcesScaceces das av deel/olded a aie PS 10
PSM CII ERE SCLIN seo e ols Faile aredsic/ cans 3) 7eievells 1» setoce: bov,s, 4.01 b s eseleiae oato ean OR ee ete 10
INGOnEOasiGal CONSIGELALIONS: vaiv ss. a/sirs.oScre's a fwd ten yee deta nme 10
Morphology of: living Polyplacophoras..5 6.0.5 6ccccecacnesences es 10
Morphology of living pleurotomarians...:......60.0s006..00sc00ecer be II
NTMSOPLeUbAate OMLOM CMY: eameiarcivercie- a o1e cicistomiotcreietete niches acetone 14
Preliminary inferences from neontological considerations.............. 16
IRECAPILULALT ON Gumres sleiaysiercrsie ole cine te ial Sloe ie one ae ele ee eno 2I
Pal eGGtOlO geal CONSIMEEATTONS © 5.54485 a.gcsva-one 0-5/0 s aes: sind video clea avery amore 22
Climpine down, the family treee cs fags sac ys snes ceive cee Seal nomle 24

The pleurotomarian-bellerophont branch to the isopleuran
Mion OplacOphonal eiyotecclceeeienveic coisas Chetan oie ool ete teens 24
aRhespolyplacophoran yoranchiycte cries c sicvsiersccieeaatvelenls ererierenels 31
Reclimbs meat neweree syste leraseee aheeie assem ienets arc eaciahe toe ete veietal teres aeotoe 33
Exploravionoar Ofer early Dranches®. co... oo 2 o< siac:eleiewiecs.01 eareroiels 34
ulbirebneatellaceae sro: ccsasicic cc's eieleaccke atcrowie ciate fe eieiatee aleve eteerettte 34
iWacluritessandeitsvalliesane sc acces coe oemineiects eet: 36
Helaprellawariauitsual esi < sieveevage.cionstciecorses crores eoreuiais ei sieternehenete 40
eA ec INOMEIAG MICATICMLTSIGUIS aie cfs chatet oie iain apace acas-@ acauarerois elcid aisveiare 0 B tavepalade Mratdiaeds 44
mppendix: Interpretation of the bellerophonts......... 06 60cicc tues «aspen om 48
RRS RCUCCSD Poa ers ectele eo eeeia nae wisisie Sarcie eT eae w wae mnie pW emaiseaine seems 55

iil

- iF eu,

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Charles D. and Marp Waux Walcott Reseach Fund

PROVUGIVE FOSSIL GASTROPODS, AND THELR
BEARING ON GASTROPOD CLASSIFICATION

By J. BROOKES KNIGHT
Research Associate in Paleontology, U. S. National Museum

(WiTH Two PratEs)

INTRODUCTION
GENERAL CONSIDERATIONS

With only one exception that comes readily to mind, the various
classifications of the class Gastropoda in current use are the work of
neontologists. The living gastropods are classified on the basis of
their morphology, largely the anatomy of the soft parts. The fossil
forms, or at least the older ones, so far as they belong to genera that
are now extinct, are given the scantest of notice and are distributed
in an almost haphazard fashion among the families erected primarily
for living forms. As neontologists have little familiarity with fossils,
unless it be the more recent ones, they are not especially struck by
the resulting incongruities. Of course they fail to take the fullest ad-
vantage of the information that the older extinct fossil forms can
furnish as to the early history of the class and its bearing on
phylogeny. Indeed the inaccuracy of such little knowledge as they
have of the more ancient fossils is apt to lead them astray.

That the work of the neontologist is nevertheless of the highest
importance is too obvious to need comment. He has the entire animal
available to him, including the soft parts, and in the main he has
made much of his opportunities.

The paleontologist, on his part, suffers from the severe handicap
that he can never observe directly the soft parts of the forms that
he studies. In a sense he is forced into the role of a mere concholo-
gist. Unfortunately, many paleontologists, inadequately trained in
zoology, surrender with resignation, if not with complacency, to what
appear to be the necessities of the situation. Nevertheless, it is pos-
sible to infer from fossil shells somewhat more of the probable gen-
eral anatomy of the soft parts than is commonly done and these

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 13

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

inferences, if made with due caution, can be useful. Of course they
do not stand on the same plane as direct observation; nevertheless
to neglect them, or to refuse to give them recognition, however
guarded, would be unscientific indeed.

The exception to my original statement that the classification of
the Gastropoda is largely the work of neontologists is the work of
Wenz, begun in 1938 and unhappily interrupted by his death soon
after the close of the second World War (Wenz, 1938-1944). He
lived to complete only that part dealing with the prosobranchs. Wenz
was a paleontologist with an excellent training. He acquired some
familiarity with the older fossil gastropods as a pupil of Prof. Em-
manuel Kayser and especially of Prof. Ernst Koken, of Tubingen.
His field of specialization since his student days was Cenozoic non-
marine gastropods, a field that did not qualify him particularly for
the task he undertook. The novelties introduced by Wenz in 1938
into the classification of the Gastropoda were not in the highest cate-
gories but at the familial level. He made a distinct contribution in
erecting many families, subfamilies, and superfamilies for extinct
genera for which there had long been a need, but the inherent diffi-
culties of working with skeletal material alone and his relative un-
familiarity with the older marine forms made many of his new fami-
lies mixtures of incongruous elements, and their placement in the
higher categories is not always fortunate.

Perhaps the outstanding contribution of Wenz’s work in 1938 to
the fundamentals of gastropod classification was his suggestion that
the isolated, symmetrically paired dorsal muscle scars of Tryblidiwm
(Tryblidiacea) might be a very primitive character suggesting the
segmentation of the chitons (Wenz, 1938, p. 59). However, in 1938
he allowed himself to be influenced by this idea in constructing his
taxonomic hierarchy only to the extent of erecting a separate super-
family, Tryblidiacea, for the genera with symmetrically paired dorsal
muscle scars instead of including them with the superficially similar
Patellacea, as had been done in effect by previous workers. As is
generally recognized, the symmetry in the Patellacea is secondary and
superficial, not primitive.

Two years later Wenz proposed a more radically revised classi-
fication of the major categories of the Gastropoda (Wenz, 1940).
He recognized a major dichotomy within the Gastropoda (excluding
the Loricata) between what he regarded as two subclasses, the
Amphigastropoda (bilaterally symmetrical, primitively orthoneurous,
with a saucer-shaped, conical, or symmetrically spiral shell) and the
Prosobranchia (asymmetrical, chiastoneurous, with asymmetrically

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 3

coiled shell). He elaborates somewhat his invaluable earlier views on
the tryblidians but he does not follow the logic of his position and clas-
sify them with the chitons. Instead, because of the discovery of multi-
ple paired dorsal muscle scars in the supposed bellerophont Cyrtonella,
he classifies the bellerophonts with the tryblidians in a subclass, the
Amphigastropoda. This action I do not regard as well taken (Knight,
1947b, p. 264, and appendix to this work). Naef, a neontologist, had
made a somewhat similar division at an earlier date with the Plano-
spiralia for the bellerophonts (he was unaware of the probable signifi-
cance of Tryblidium and its allies or possibly even of their existence)
and the Turbospiralia for the asymmetrical groups (Naef, I9I1,
p. 159). Naef’s Planospiralia, unlike Wenz’s Amphigastropoda, was
looked on as streptoneurous and, of course, prosobranch.

In the interval between the first draft of the present paper and its
completion, an interval required for the preparation of drawings, a
significant paper on the aspidobranch Gastropoda and their evolution
appeared. This paper, by the distinguished anatomist and physiolo-
gist, C. M. Yonge (1947), reports the results of some revealing in-
vestigations on the anatomy and functioning of the pallial organs of
some aspidobranchs. Yonge does not stop with the recording of ob-
servations but proceeds to apply his findings to an interpretation of
gastropod evolution just as I have done from a different set of obser-
vations. Both Yonge and I have accepted certain findings and inter-
pretations from previous workers and to that extent have a common
background. Hence it is not surprising that there is much basically
the same in each interpretation. On the whole our acceptance or re-
jection of the suggestions of previous workers is gratifyingly similar.
A minor difference is that he regards Wenz’s suggestion that the
tryblidians are pretorsional gastropods only as possible (Yonge, 1947,
p. 485). With some rearrangements and differences in emphasis from
Wenz I accept this as probable. Yonge regards the bellerophonts as
prosobranchs, just as I do, and thus rejects Wenz’s view that they
were “primitively orthoneurous.” However, he appears to harbor an
unexplained and undocumented idea that although they are symmet-
rical prosobranchs they had a single dorsal and median retractor
muscle (Yonge, 1947, p. 490, fig. 31a). It is my view that the bel-
lerophonts are prosobranch gastropods that have undergone torsion
and have retained a high degree of primitive bilateral symmetry in-
cluding a single symmetrical pair of retractor muscles attached at the
distal ends of the columella (Knight, 1947b).

Yonge proposes some phylogenies (Yonge, 1947, p. 490, fig. 31a)
toward which I am compelled to be skeptical. I am skeptical of the

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

supposed origin of the Neritacea and the pectinibranchs as branches
arising independently and directly from the bellerophonts. The great
expansion of the pleurotomarians in the Paleozoic when they over-
shadowed all other contemporaneous gastropods in diversity of form
and number of genera and species provided possibilities that cannot
be neglected. For example, the asymmetrical neritaceans and the
pectinibranchs may have been derived from asymmetrical ancestors
such as some of the numerous and varied pleurotomarians. The
pleurotomarians (Pleurotomariacea) include much besides “Pleuro-
tomaria.” That the present-day Theodoxus and the pectinibranchs
are not derived from the present-day “Pleurotomaria” one can readily
concede. It is equally unlikely that the present-day Haliotis, Scis-
surella, the fissurellids, or the patellids are correctly derived from the
present-day “Pleurotomaria,” as Yonge seems to imply. I am not even
prepared to accept Mikadotrochus beyrichi (Hilgendorf), the species
from which Yonge derives most of his ideas of pleurotomarian anat-
omy, as properly referred to the genus Pleurotomaria Sowerby. That
all these may have had common ancestors more advanced than
bellerophonts, i. e., advanced to the pleurotomarian stage or farther,
seems probable.

The classifications of neontologists are based actually on com-
parative anatomy, that is to say on morphology, from which they
attempt to infer phylogeny, but phylogeny, or descent with change in
time, is held very much in the background as an ideal only. The pa-
leontologist alone has spread before him the time sequence, the order
in which forms appeared in time. This has been called chronogenesis.
Chronogenesis is not a perfect tool, for the fossil record is far from
complete and the recognition of phylogenies involves supposed relation-
ships inferred from imperfectly known morphological criteria. Never-
theless, it is a useful tool, if used with caution, and is becoming more
and more useful as our knowledge of the life of the past increases.

In summary, all classifications are provisional and hypothetical,
based on inferences from more or less complete observations of vari-
ous phenomena. Certainly any classification based wholly on neon-
tological data or with inadequate attention to or understanding of
paleontological data must be almost as defective in the very nature
of the case as would be the converse. The present classification is
offered as one that at least attempts to give full weight to paleon-
tological data and their bearing on phylogeny. It is admittedly pro-
visional.,

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 5

PROPOSED CLASSIFICATION

Changes in classification.—The principal novelties of the classifica-
tion given below consist of the following:

The Polyplacophora are returned to the Gastropoda as one order
of a subclass, the Isopleura, proposed by Lankester in 1883. The
order Monoplacophora (Tryblidium and its allies) is added to the
Polyplacophora. I am allowing the Aplacophora to stand close to the
Polyplacophora, as do most authors, although without strong con-
viction. They do not occur as fossils, and paleontology has no
light to throw on them. They are probably degenerate, not primitive.
These three orders will make up the Isopleura.

In the subclass Anisopleura, also proposed by Lankester in 1883, are
included as superorders the Prosobranchia, Opisthobranchia, and Pul-
monata, while to the primitive prosobranch superfamilies Bellero-
phontacea and Pleurotomariacea of the order Archaeogastropoda is
added a third, the Macluritacea. Except as affected by the foregoing
the remaining more advanced archaeogastropod superfamilies are left
untouched as Wenz left them, not because Wenz’s treatment is satis-
factory but because a complete revision is beyond the scope of this
paper. Such a revision is well under way, however, and perhaps in the
not too distant future the results may be published.

Phylum Mollusca Cuvier
Class Gastropoda Cuvier
Subclass Isopleura Lankester
Order Monoplacophora Wenz 1
Order Polyplacophora Blainville
Order Aplacophora Jhering
Subclass Anisopleura Lankester
Superorder Prosobranchia Milne-Edwards
Order Archaeogastropoda Thiele
Superfamily Bellerophontacea Ulrich and Scofield 2
Superfamily Pleurotomariacea Wenz ?
Superfamily Macluritacea Gill 1

Other archaeogastropod superfamilies are not considered here, nor
are the following orders and superorders:

Order Mesogastropoda Thiele
Order Neogastropoda Wenz
Superorder Opisthobranchia Milne-Edwards
Superorder Pulmonata Cuvier
Incertae sedis. Pelagiella Matthew, 1895, and allies. Possibly not
gastropods.

1 Considered in some detail.
2Only the earlier, more primitive genera and some living ones considered.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 7

Chronogenesis and range in time—In order to give an over-all
view of the range in time of the two orders of the Isopleura and the
three most primitive superfamilies of the anisopleuran prosobranchs,
a diagram is presented (fig. 1). It will be noted that the major
dichotomy in time (as well as in morphology) is between the Iso-
pleura and Anisopleura in the early Cambrian, at the beginning of
the fossil record. It will be noted also that two of the three primi-
tive superfamilies of the prosobranch Archaeogastropoda, the Mac-
luritacea and Bellerophontacea, have been extinct since Devonian and
Triassic times, respectively. Only the long-ranging Pleurotomariacea,
late Upper Cambrian to Recent, has carried through in several special-
ized relic families. These give us some clue to the morphology of the
soft parts and to the physiology and embryology of the primitive
prosobranchs,

EXPLANATORY NOTES

Technical terms.—As far as possible the use of technical terms
(other than the formal scientific names of systematic categories) has
been avoided. With exceptions to be noted the morphological
terms employed are so much in general use by both paleontologists
and malacologists that it seems unnecessary to define them.

The term “hyperstrophic” is not a new one but experience suggests
that many have only a hazy idea of its meaning. It refers to that

Ficure 1.—Range in time of the more primitive categories.

For the benefit of the neontologist interested in geologic time in terms of years
and unfamiliar with recent work, the following data are arranged from the
Report of the Committee on the Measurement of Geologic Time of the Division
of Geology and Geography, National Research Council, for 1949-1950 (p. 18):

Beginning in

approximate Approximate
number of ength in
millions of millions of
Period years ago years

UAE rh atv te cistele ise sic iorcvereuelelaleveeta, systole I I
ENtIAGY ete eh alot toate see eve atavstonsvareia sia 60 59
IGrefaccotismeeciie craters nisicralcinle <olatotareiaravata oT 130 70
VME Aogsbecouds Hose UOL OME UDOBDOC 155 25
RUE LASGUC MEE oe elexevayateverateley sxe fieke..ev erate) eye inv eic'e)s 185 30
RETA IAT I tepeclapeyciotetttstaiels eteisisteley sve: sve sicisiero'e 210 25
(Chmethisdae) aon HOAUOude DAOOD MO DUO OL 265 55
LDYSerhebe, croc COMPOS OSHA BD GUDIDICS On ONUODG 320 55
(SENET e) eae Kin BECHER ReO IDO ROGER IT DOORS 360 40
Weclavicianiee cris ciciehisisscvclavcla sini sle miviaiainye rete 440 80
(Chin sent aGag etoile DAD ne DOORS Dov oODOS 520 80

Computed probable errors in beginnings: Quaternary + 50 thousand years,
Tertiary + 1 to 2 million years, Mesozoic + 5 million years, and Paleozoic + Io
million years.

b-f,

ef,

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, II7

FIcurE 2

Isotrophic coiling. Symmetrically coiled in a plane with the sides mirror
images of each other. The example is a diagrammatic restoration of a gen-
eralized bellerophont. Note the symmetrical and paired ctenidia, the rectum
passing through the pericardium and terminating in the anus between the
ctenidia and close to the slit, and the symmetrical and paired auricles of the
heart.

Asymmetrical coiling. b, Orthostrophic coiling in the pleurotomarians. Al-
though the shell is asymmetrical and orthostrophic the rectum still passes
through the pericardium and terminates between the ctenidia close to the
slit. Many of the organs such as ctenidia, auricles, etc. are paired. The
diagram shows a dextral pleurotomarian. No certainly sinistral ones are
known. c,d, Orthostrophic coiling at an advanced stage of asymmetry. In
the dextral forms the right ctenidium and auricle are lost, the rectum has
moved to the right and no longer passes through the pericardium, and the
slit has disappeared. c shows a sinistral orthostrophic gastropod and d a
dextral one. Note that the arrangement of the shell and the internal organs
in each is the mirror image of the other. A tremendous majority of living
gastropods are dextral orthostrophic.

Hyperstrophic coiling. e, sinistral and f dextral. Comparing the dextral
and sinistral hyperstrophic forms with their orthostrophic counterparts, the
relative positions of the corresponding internal organs are the same in each
but the direction of asymmetry of the shell is reversed. In the dextral
hyperstrophic form the spire protrudes to the left side instead of to the
right. The internal organization is dextral, but the shell if oriented in the

[legend continued on opposite page]

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 9

sort of coiling in which the shell is inverted and what appears to be
the spire is homologous with the base of orthostrophic forms. It is
as though the normal spire were pushed through, protruding on the
side that is normally the base and the side that normally has the spire
resembles a base. The shell resembles superficially a sinistral shell
but the soft parts are dextral. A hyperstrophic sinistral shell re-
sembles a dextral one but the soft parts are sinistral.

The term “orthostrophic” is employed for the normal coiling of
the great majority of asymmetrical gastropods both dextral and sinis-
tral. The true sinistral gastropod is in all respects a mirror image of
a dextral gastropod.

The term “isostrophic” is introduced as an adjective to describe
the sort of coiling that is found in many nautiloid and ammonoid
cephalopods, and particularly in the bellerophont gastropods. It may
be exogastric as in the cephalopods or endogastric as in the gastro-
pods. Isostrophic coiling is symmetrical with the left and right sides
mirror images of each other.

Text figure 2 illustrates the different types of coiling described
above.

The term “Cambrian” is employed here in the current American
sense (Howell et al., 1944, pp. 993-1004) in which beds of Trema-
docian age are excluded from the Cambrian. Those are placed as
late Lower Ordovician. It is important that the European reader bear
this in mind.

Illustrations—In addition to certain diagrammatic drawings to
illustrate various points under discussion I have included drawings of
generalized restorations of a number of characteristic Paleozoic gen-
era mentioned in the text. Many of these are yet unfamiliar to any
but specialists and it is hoped that the drawings will be of assistance
to the general reader in visualizing what must be unfamiliar genera
to many. Although these were/ made from actual specimens of spe-
cies, they are restorations intended to illustrate generic characters
and are not accurate enough to be used for the identification of
species.

References.—The list of references will be found on pages 55 to
56. In the text, references to the list are cited in parentheses by
author and date. Since I published some years ago descriptions and

customary way with the spire upward appears to be sinistral. In this paper
all illustrated species judged to be hyperstrophic are oriented with the spire
downward for this brings the aperture to the same side as in a conven-
tionally oriented orthostrophic shell. See plate 2 on which the Pleuroto-
mariacea shown are dextral orthostrophic and the Macluritacea are dextral
hyperstrophic. ve nS

To SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 117

figures of the type species of all names of genera based on Paleozoic
species published before 1938 (Knight, 1941), no further references
to such genera will be given here. References to Paleozoic genera pub-
lished since 1937 appear in the list. For post-Paleozoic genera the
reader is referred to Wenz, 1938-1944, which will suffice for many
purposes. If this rather rare work is unavailable, many works on
conchology or malacology will do.

ACKNOWLEDGMENTS

I am indebted to a number of colleagues who have read my manu-
script at one stage or another of its development. Among these are
Dr. G. Arthur Cooper, Dr. H. A. Rehder, and R. Tucker Abbott, of
the U. S. National Museum, and William T. Clench, of Harvard
University. Perhaps I am indebted most to Prof. Raymond C.
Moore, of the University of Kansas, and to Dr. L. R. Cox, of the
British Museum (Natural History). The former has lent encourage-
ment over the five or six years that my ideas have been developing on
paper. Both have given the paper critical readings and have furnished
the stimulus of dissent from some of the views expressed. I alone am
responsible for departures from the orthodox.

ARGUMENT

In order to arrive at hypotheses worthy of attention one must pro-
ceed from the known to the unknown, or from the better known to
the less well known. Therefore it will be profitable to consider at
this point certain selected zoological data, well known, perhaps, to the
neontologist but relatively unfamiliar to many eee toa Later,
paleontological data will be considered.

NEONTOLOGICAL CONSIDERATIONS
MORPHOLOGY OF LIVING POLYPLACOPHORA

The chitons are regarded morphologically as the most conservative
in the basic pattern of their organization of all living gastropods, if
not of all living mollusks. In respect to certain features, the division
of the shell* into eight plates and the musculature to operate them,
they appear to be highly specialized. Likewise the remarkable shell
eyes or aesthetes appear to be developed in some genera, possibly in

8[ am regarding the polyplacophoran shell as homologous with the shells
of other mollusks. However, it should be noted that at least one recent worker
regards it as only analogous (Thiele, 1931, p. 2). Perhaps on further study
this very fundamental difference will be resolved.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT EAL

response, as it were, to the loss of the sense organs of the head.‘ It
is these specialized features, the modifications of the primitive basic
plan, that distinguish them as chitons. The basic plan of organization
is bilaterally symmetrical in all significant respects. There is a flat,
creeping foot and well-differentiated head. The head lacks the usual
sense organs, possibly due to specialized degeneration. The mouth
is anterior and the anus posterior, the digestive tract passes through
the pericardium. The gills appear to be true ctenidia and are ar-
ranged in pairs in a groove between the shell and the upper surface
of the foot on each side of the body, dominantly in the posterior part
(Yonge, 1939). The numerous paired ctenidia seem to be metameric
repetitions of a primary pair that lie on each side of the anus and
just behind the excretory pore (the postrenal gill). The heart is
dorsal and posterior. The nervous system is not twisted and shares
the bilateral symmetry of the rest of the body. Strictly speaking
there is no pallial cavity, but it seems reasonable to regard the pos-
terior and lateral parts of the groove between the shell and the foot
that contains the ctenidia and associated organs as strictly homologous
with the pallial cavity of more advanced gastropods.

There is no need here to go into a complete morphological descrip-
tion of the chitons. The features to which I wish to draw special
attention are the complete bilateral symmetry of all parts and the
posterior anus, gills, and heart. It is these features that are regarded
aS primitive and it is contended that it is the modification of these
features in the ancestral stock from which the chitons were derived
that gave rise to the Anisopleura, modification primarily through
torsion and progressively greater asymmetry.

The chitons appear first in the fossil record in late Cambrian time
and are living today. They were never abundant and for most of the
time were very rare. They have varied throughout all that vast ex-
panse of time very little indeed.

MORPHOLOGY OF LIVING PLEUROTOMARIANS

The pleurotomarians ® are classified in the subclass Anisopleura,
superorder Prosobranchia, order Archaeogastropoda. They first ap-

4 Can it be that the microscopic tubules in the shell of Tryblidium reticulatum
Lindstrom described and illustrated by Lindstrom in 1884 (p. 56) are the
tubules of aesthetes instead of some unknown parasitic organism?

5 The noncommittal vernacular name for this group is employed at this point
in its broadest sense, as an informal synonym of Pleurotomariacea. The group
has been treated at one time or another as a genus, as a family, or as a super-
family. Wenz in 1938 assigned a little over 200 genera and subgenera, fossil
and living, to the Pleurotomariacea.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

pear in the fossil record in late Upper Cambrian time.* They were
the most numerous, varied, and abundant of all gastropods through-
out succeeding Paleozoic time. They continue in diminishing num-
bers and variety through the Mesozoic and carry through to the
present as a few genera, in a few families, the most abundant and
diversified of which represent two late specializations for rock cling-
ing (Haliotidae, Fissurellidae). Another living family, composed of
rare and very tiny forms, is the Scissurellidae. The fourth family
of living pleurotomarians, the Pleurotomariidae, is represented in
present seas by four rarely seen but large and handsome deep-water
species, of great morphological interest because they are seemingly
little-changed descendants of early and primitive anisopleuran gastro-
pods. Although entire specimens are very rare, there have been a
number of successful dissections with which are associated the names
of W. H. Dall, E. L. Bouvier and H. Fischer, and of M. F. Wood-
ward. Dissections have been made also of some of the abundant but
specialized Haliotidae, Fissurellidae, and Scissurellidae, but the Pleu-
rotomariidae appear to be less conspicuously specialized for particular
environments and therefore more significant for the present purpose.*

This is no place to consider the minor anatomical details of the
pleurotomarians but certain major features are of importance for our
purpose. First, in common with all anisopleurans in which the fea-
tures are not obscured by later developments, all display the effects
of torsion in that the primitively posterior anus and pallial complex

6 The genera I refer to, four in number, include three with a deep U-shaped
or V-shaped sinus in the outer lip regarded by most paleontologists as homol-
ogous to a slit. These are Sinuopea Ulrich, 1911 (pl. 2, fig. 1), Schizopea Butts,
1926 (pl. 2, fig. 2), and Dirhachopea Ulrich and Bridge, 1931. The fourth,
Taeniospira Ulrich and Bridge, 1931, has a moderately deep pleurotomarian slit
and a slit band.

7It will be well here to point out that primitive prosobranch gastropods have
not a single retractor muscle but a pair of retractor muscles. For example, the
living representatives of two of the four existing families of the Pleuro-
tomarianea, the Scissurellidae and the Haliotidae, have a pair of shell or retrac-
tor muscles, and a third, the Fissurellidae, has a crescentic muscle accepted as
compounded from an original pair. In the Bellerophontacea, supposedly the
immediate forerunners of the pleurotomarians, there is likewise a single pair;
and in the Neritacea, seemingly an ancient branch from the pleurotomarian stock,
and the very primitive Macluritidae there is also a pair. In the living repre-
sentatives of the Pleurotomariidae alone, of the supposedly primitive stocks,
is there a single retractor muscle. This suggests strongly that in this respect
these have lost one of the primitive muscles and have advanced far toward
Calliostoma Swainson, 1840, in the Trochidae, to which they may be more
closely related than to the more primitive pleurotomarians.

NOES PRIMITIVE FOSSIL GASTROPODS-——-KNIGHT 13

are found in an anterior position above the head as though they had
been twisted into that position. All have a helicoidally coiled, asym-
metrical shell at least in late larval stages. But in spite of torsional
asymmetry and the beginnings of lateral asymmetry they retain, as a
primitive character fully retained in no other group of living ani-
sopleuran gastropods, paired visceral organs, including paired ctenidia,
paired auricles of the heart, paired kidneys, etc. The digestive tract
passes through the pericardium and the anus discharges between the
two paired ctenidia. These are primitive characters and they remind
one strongly of the bilaterally symmetrical pairing of the homologous
organs in the isopleuran Polyplacophora. They suggest that the Ani-
sopleura were derived ultimately from bilaterally symmetrical, iso-
pleuran ancestors.

Recent and fossil pleurotomarians always, or nearly always, show
one distinctive shell feature by which they may be recognized almost
at a glance. This is an emargination in the outer lip of the shell. In
some of the earliest species it takes the form of a rather deep U-shaped
or V-shaped sinus. In others the sinus is V-shaped and it may culmi-
nate in a short slit or notch. Still later appear forms with a deep slit
and still other modifications, such as a row of tremata, developed in-
dependently in several genera, or the apical hole in the shell of the
typical fissurellids. In all living pleurotomarians that have been ex-
amined the discharge end of the anal tube lies close to the apex of the
emargination. Yonge has shown from studies of living examples of
Haliotis Linné, 1758 (Yonge, 1947, p. 449), of the anatomy of a pre-
served specimen of Mikadotrochus beyrichi (Hilgendorf) (op. cit., p.
454), and of the described anatomy of /ncissura lytteltonensis Smith
(op. cit., pp. 449-458), as well as of living examples of the more highly
specialized Fissurellidae, that the respiratory current is created by cilia
on the filaments of the ctenidia. The water is drawn into the mantle
cavity above and on both sides of the head. It passes backward beneath
the ctenidia, impinging on the osphradia enroute, then upward between
the ctenidial filaments and outward through the slit or its equivalent.
Close to the inner end of the slit (or row of tremata) is the anus at
the distal end of the anal tube. The currents, possibly aided by con-
tractions, carry the faeces out through the slit. This is a highly im-
portant matter of sanitation for any gastropod with the anus within
the mantle cavity and directed anteriorly so as to discharge between
a pair of ctenidia. Obviously such a mechanism would not be impor-
tant for forms with a posterior anus, such as isopleurans, or for
those such as the more advanced anisopleurans with an anterior anus
but with only one ctenidium on the upstream side, as it were, of the

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

ciliary currents passing through the mantle cavity, or secondarily
with a more or less posterior anus as in the opisthobranchs.

ANISOPLEURAN ONTOGENY

There are inherent technical difficulties in the rearing and studying
of such extremely small and delicate organisms as the early embryos
of primitive gastropods. Because of the complexity of the transfor-
mations and the confusing differences in detail from one species to
another it is difficult to make generalizations in terms that will be
valid in detail for even the few forms for which much is accurately
known of the early ontogeny. Furthermore it is difficult to avoid
attributing to the embryo adult anatomical features which occur only
as rudiments, often as only a few cells not obviously organized, if
present at all in the embryo. For example, it is commonly thought
that in the process of torsion the gastropod becomes so twisted that
the pallial cavity with the pallial complex including anus, ctenidia,
kidneys, heart, etc., is translated bodily from a posterior position to
an anterior position above the head. In effect this is true but in detail
it is not, for in such primitive genera as Haliotis and Patella Linné,
1758, for example, the “proctodaeum and the solid mesoderm rudi-
ments of the kidneys are the only representatives of the pallial com-
plex when torsion begins. Even in Viviparus Montfort, 1810,® in
which the developmental stages are abbreviated owing to viviparity,
the single ctenidium and the visceral part of the pleuro-visceral loop
do not develop until . . . after torsion is complete” (Crofts, 1937, pp.
262-263). However, in spite of these difficulties a significant series
of events does occur in a definite order.

The early trochophore larva has a dorsal shell gland and a stomo-
daeum (rudimentary mouth) situated immediately below the ciliated
ring of the velum on the ventral side. The shell, secreted by the shell
gland, develops from a small disc to a rather deep cup containing the
dorsal hump. The proctodaeum (rudimentary anus) is moved ven-
trally toward the stomodaeum. In the process the rudimentary gut,
still without open mouth or anus, is bent into a rough U-shape. This
operation is called flexure and is regarded as distinct from torsion for
which, however, it lays the foundation. The rudimentary foot appears
between the stomodaeum and proctodaeum. The pallial cavity appears
as an invagination posterior to the foot. In the meantime the shell

8 For Paludina, the name employed by Crofts and other embryologists, I am
substituting the name Viviparus, today regarded as the correct name of the genus
concerned.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 15

has continued to grow and, owing to secretion of shell matter more
rapidly on the posterior margin, it takes on an exogastric roughly
nautiloid form with the primitive apex directed forward.

A highly significant organ, the development of which is completed at
the end of the pretorsional stage, is the single “velum retractor mus-
cle” first carefully studied by Crofts in Haliotis (Crofts, 1937). The
muscle before torsion is asymmetrically placed and slightly spiral in
such a way that its retraction rotates the dorsal hump in a counter-
clockwise direction when viewed dorsally. In passing it will be ad-
vantageous to note that the velum retractor persists through life in
Haliotis as the small left-hand shell muscle and that the hypertrophied
right-hand shell muscle, homologous with the single columellar or re-
tractor muscle in most gastropods, is not at this stage represented by
a recognizable rudiment of even a single cell. It is probable that the
left-hand retractor muscle in the adults of the more primitive aspido-
branchs is entirely homologous with that of Haliotis. In more ad-
vanced types it is lost before maturity.

In Haliotis torsion begins at about 30 hours after fertilization of
the egg. Crofts (1937, pp. 233-234) reports that the first 90° of
torsion takes place in 3 to 6 hours as a result of contraction of the
“velvum retractor muscle.” The full 180° torsion is not completed
until 8 or 10 days later and apparently results from differential
growth.

There are curious differences in both the process and time of tor-
sion as reported by different authors for different species and even
for the same species. Some of these differences may be caused by
the difficulties in observing accurately such small and refractory sub-
jects, but most of them probably reflect actual differences between
species. Nevertheless there is general agreement on the fact of 180°
torsion at an early embryonic stage.

The torsion results in the pallial cavity’s moving from a posterior
to an anterior position relative to the foot. Although the organs of
the pallial cavity have not yet appeared when torsion begins or are
extremely rudimentary they eventually mature after torsion in an
anterior position even though their primitive position must have been
posterior. Likewise the commissures of the visceral nerve complex
mature after torsion as though they had been crossed to a figure 8
during the process, although during torsion they were far too short
and rudimentary to be crossed. Torsion, of course, affects relations
of the shell to the head and foot so that its apex points to the rear
of the head instead of forward.

Before torsion there is some asymmetry in one respect or another,

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

in part no doubt anticipatory in nature and chargeable to accelera-
tion, but after torsion asymmetry develops apace. It is only less
marked in those forms that develop primitive paired organs than in
those that develop only one member of the primitive pair, usually
the definitive left member. The shell is no longer approximately
bellerophontiform, but coils in a laterally asymmetrical, helicoid spiral
with the spire pointing backward. That in certain groups the shell
then becomes symmetrical (Diodora Gray, 1821, for example) or
that secondary detorsion occurs (opisthobranchs) with a high degree
of superficial secondary symmetry is irrelevant to our present dis-
cussion. Nor is it relevant that in a few forms the torsion is clock-
wise resulting in sinistrality (see fig. 3).

To recapitulate, the anisopleuran veliger larva is provided with a
dorsal shell gland ; the gland secretes a shell that grows by marginal
accretion and soon becomes cuplike; concurrently the pallial cavity
is invaginated and the body, with the rudimentary alimentary canal,
is bent to a U-shape with the procotodaeum within the pallial cavity
posterior to the stomodaeum and separated from it only by the rudi-
mentary foot. The U-shaped bending constitutes flexure. The next
step is torsion by which the dorsal hump with the pallial cavity is
twisted 180° in a counterclockwise direction (as seen from above)
relative to the foot, thus laying the foundation for the prosobranch
and streptoneurous conditions. Next comes, as a separate step, the
development of lateral asymmetry and the helicoid spire. There are
in some advanced stocks still further developments, including detor-
sion which brings about the opisthobranch condition, euthyneury, and
in extreme cases secondary symmetry of a high order.

PRELIMINARY INFERENCES FROM NEONTOLOGICAL
CONSIDERATIONS

From anatomy.—It is inconceivable that the living anisopleuran
gastropods, which show torsional and generally lateral asymmetry
and which are members of the Mollusca, a phylum characterized by
basic bilateral symmetry, can be at all primitive in respect to those
features. The most ancient anisopleuran group with living represen-
tatives, the Pleurotomariacea, appears first in the late Cambrian.
Living pleurotomarians show vestiges of bilateral symmetry in the
retention of paired visceral organs along with full torsional asym-
metry and laterally asymmetrical coiling. The Polyplacophora living
today are equally as ancient as the pleurotomarians. They are ob-
viously specialized in respect to the eight-pieced shell, but they retain

NOP TS PRIMITIVE FOSSIL GASTROPODS—KNIGHT 17

FIGURE 3

Torsion in the embryo of Viviparus viviparus (Linné). It should be noted

that owing to the viviparity the developmental stages in Viviparus are ab-
breviated. Hence for this reason and because of the highly diagrammatic
nature of the drawings (from Naef, 1011, fig. 8, in part) the picture pre-
sented is somewhat oversimplified. It is all the more comprehensible for those
reasons.

Stage where flexure is in progress but torsion not begun. To the left of the
figure is the ciliated velum, the cup-shaped shell is above and the rudi-
mentary foot below. The digestive tract is dotted with the mouth below
and to the left and the anus high and to the right of the figure.

The beginning of torsion. The mantle cavity has appeared and with the
anus is turned a little to the right of the animal.

Torsion a little more than halfway completed. The anus and mantle cavity
are now to the right and a little to the front.

Torsion completed. Note that the mantle cavity with the anus is now in
front and above the head, it final position. The shell has become
bellerophontiform.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 117

strict bilateral symmetry. They cannot be ancestral to the pleuroto-
marians, but they very plausibly point the way to that more remote
ancestor of both chitons and pleurotomarians which must be looked
for first in Lower or Middle Cambrian rocks unless it became extinct
before Cambrian time with its record irretrievable. Several very
distinguished neontologists have speculated as to the probable nature
of this common ancestor of both and, indeed, of all the Mollusca.
The usual conclusion is that it was a mollusk with a single, low,
conical shell, bilaterally symmetrical in all respects, with the anus and
pallial complex in the rear, with a differentiated head and a flat creep-
ing foot. In epitome, it would have the basic bilateral symmetry of

FIGURE 4

Scheme of a hypothetical primitive mollusk viewed from the left side. a, anus;
c, g, cerebral ganglion; f, foot; g, gill, in the pallial cavity; go, gonad; h,
heart; k, kidney; Jac, labial commissure; m, mouth; pa, mantle; pan, pallial
nerve; fe, pericardium; pg, pedal ganglion; plg, pleural ganglion; ra, radula;
rpo, renopericardial orifice; st, stomach; stg, stomatogastric ganglion; vg,
visceral ganglion. (After Pelseneer. )

the Polyplacophora but with a single shell, as in the Anisopleura, but
neither coiled nor asymmetrical. Figure 4 shows a reconstruction of
such a hypothetical ancestral gastropod, a reconstruction based on
pure deduction before anyone had suspected the possibility that sup-
posed Cambrian capulids or patellids that we now recognize as the
Monoplacophora had just about the same anatomy. Figure 5 shows
a restoration of a generalized monoplacophoran.

From ontogeny.—Before torsion the cup of the larval shell deepens
with flexure of the intestinal tract and because of more rapid growth
at the posterior margin takes the form of the beginning of an iso-
strophic or bellerophonlike coil but with the apex or rudimentary coil
forward. It seems reasonable to suppose then that the descendants
of our hypothetical ancestral gastropod may have passed through
similar stages in the initial process of becoming coiled. Indeed coil-

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 19g

ing could hardly have occurred in any other way. The development
of a higher and higher shell and the initiation of coiling symmetri-
cally in a plane are processes that lend themselves to gradual evolu-
tionary development. If the fossil record is sufficiently complete, we

Pia

Stomach-—=y- - >

Muscle._ #
scars ¥

FIGURE 5

Schematic restoration of a generalized scenellid treated as though it were trans-
parent. In making the restoration there were employed the concepts of un-
torted bilateral symmetry suggested by the muscle scars of Archaepliala.
a, Left side view; b, from above. Except for the muscle scars, note the
resemblance to the hypothetical primitive mollusk (fig. 4). The latter was
suggested by Pelseneer without reference to scenellids which he regarded as
anisopleurans (i .e., Docoglossa).

should expect to find among the earliest gastropods forms with
complete bilateral symmetry and a low, cup-shaped shell with an apex
somewhat in front of the center, others that maintain the bilateral
symmetry with a higher, narrower shell and the apex partly coiled
forward, and still others with a complete coil, all steps necessary to a
gradual evolution. As will be shown in the following parts of this

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

paper, that is precisely what we do seem to find in the earliest fos-
siliferous rocks.

The next ontogenetic step, the sudden torsional twisting, is spec-
tacular and of the highest significance. Since torsion is not a phe-
nomenon that lends itself to gradual step-by-step development it is
highly probable that it occurred just as suddenly phylogenetically as
it does today ontogenetically. It is possible, if not probable, that
torsion originated as the result of a genetic mutation having its
phenotypic expression effective at the veliger stage of the ontogeny
(Garstang, 1929, p. 89). This is the view that was set forth by W.
Garstang and that has radically altered the thinking of many students
of the Gastropoda. If Garstang’s view is true, an isopleuran parent
may have produced anisopleuran offspring. What could only be
regarded as a monstrosity if it had gone no farther was so success-
ful that the strain that carried the genes as a part of its heritage
prevailed in competition and eventually brought into being an entire
new subclass. (Class in accordance with the usual classification.)

If torsion did arise suddenly in some such manner as Garstang postu-
lated, then the adults of the first torted stock should have resembled
their parent in every respect except that they had undergone torsion
as larvae. They would have retained all their paired organs sym-
metrically developed and their shells would have retained their sym-
metry but with the apex or coil now in a posterior position. They
would have retained other peculiarities of the parent stock such as
the basic plan of ornamentation. The anus and pallial complex, how-
ever, would be above the head and directed forward because of tor-
sion, and since the immediate parent with a posteriorly directed anus
and pallial complex had and needed no special provision for clearing
the pallial cavity of waste products, the newly torted offspring would
be like the parent in this respect. That is to say, it would have no
anal emargination in the lip of the shell. In the Early and Middle
Cambrian are shells that seem to meet these specifications.

Since the newly acquired orientation results in a position of the
anus and pallial complex that would seem to make it difficult to avoid
fouling the ctenidia with waste products, we might expect that muta-
tions providing a mechanism for ready disposal of the faeces and
urine without fouling would have survival value. Hence it is not
surprising to find in Upper Cambrian rocks the first bellerophonts
with an anal emargination. It is then present in three bellerophont
families. The forms without this sanitary provision disappear shortly
thereafter.

NOS, £3 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 21

The significant embryological studies of neontologists were made on
asymmetrical anisopleuran gastropods, the asymmetrical development
following closely on torsion. Consequently we should not be sur-
prised to find that lateral asymmetry appeared in the paleontological
record soon after the establishment of a line of isostrophic gastropods
(bellerophonts) with only torsional asymmetry. This expectation is
realized in the appearance of the first known pleurotomarians in late
Upper Cambrian rocks.

RECAPITULATION

Summarizing our inferences from neontological data we arrive at
the following hypotheses which may be tested against paleontological
data. The first is that the Polyplacophora and the pleurotomarians
were derived from a common ancestor with complete bilateral sym-
metry. We infer also that the Polyplacophora have evolved from that
common ancestor through the segmentation of the shell but retention
of bilateral symmetry. We may further infer that the pleurotomarians
have evolved first through the introduction of torsional asymmetry
by a single mutation phenotypically effective at the veliger stage of
ontogeny (bellerophonts) and later through the initiation of the
lateral asymmetry that characterizes all the Anisopleura other than
the bellerophonts. (See fig. 6.) Lateral asymmetry is carried pro-
gressively much farther in more advanced groups. We may still
further infer something of the probable characters of the isopleuran
common ancestor of the Polyplacophora and the pleurotomarians and
of the intermediate stages between the pleurotomarians and that
ancestor.

The immediate predecessor of the pleurotomarians should have had
all the characteristics of that group except lateral asymmetry. It
should have had torsional asymmetry but lateral symmetry ; it should
have been coiled tightly or loosely or with a curved apex pointing to
the rear in a plane, with each side the mirror image of the other
(isostrophic). It should have had an anal emargination when fully
established but not when it first suddenly came into being. It, in turn,
should have had an immediate predecessor with a high, conical shell
with curved apex as the first step toward isostrophic coiling. The
high conical shell would of course have been deep and would have so
crowded the multiple paired pedal muscles that there would have been
room for only a few, perhaps only a single pair. If torsion had taken
place the apex of the shell would have pointed backward, if not it
would have pointed forward. The last-mentioned stage should have
had as a predecessor an untorted mollusk with complete bilateral

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

symmetry and a low, shallow conical shell with little or no flexure of
the viscera and, of course, no torsion.

Since, by hypothesis, this most remote stage was ancestral to the
Polyplacophora as well as to the Anisopleura it might conceivably
display characters basic to the transverse segmentation of the shell
into separate plates, characters such as multiple transversely paired

Waa

~-L Le

FIGURE 6

a, Schematic drawing of a primitive pleurotomarian seen from above (modified
from Naef, 1or1). The primitive anal emargination is shown as a
U-shaped sinus. The anal tube leading from the stomach is shown as
passing through the pericardium and terminating close to the emargina-
tion and between a pair of ctenidia.

b, Similar drawing of a primitive sinuitid bellerophont (also modified from
Naef, 1911). It is thought to have been very like the primitive pleuro-
tomarians but with complete lateral symmetry. Both have undergone tor-
sion and the pallial complex is anterior instead of posterior.

pedal or shell muscles. It might have possessed in a rudimentary form
traces of the tubules that carry the aesthetes in the tegumentum of
modern Polyplacophora.

PALEONTOLOGICAL CONSIDERATIONS

In order to dispel certain misconceptions widely prevalent in neon-
tological circles and, alas, occasionally met with in paleontological
circles, a few words in general terms about the gastropods of the
Cambrian period may be helpful. These misconceptions arose largely
through the efforts of paleontologists of an earlier day to. place spe-
cies, inadequately understood because of poor preservation or some
other cause, in established genera, often in order to avoid erecting

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 23

new genera for them. Since much of the evidence is not explicitly
in the literature and cannot be introduced here without making this
paper too diffuse, I am forced to speak dogmatically on some points.

There are no Platyceratidae known from rocks older than Middle
Ordovician, and no Capulidae in the Paleozoic. That there are both,
is a common error. Pleurotomarians do not occur throughout the
Cambrian section so far as is known, and only the Late Cambrian
bellerophonts are readily recognized as such. There are only four
pleurotomarian genera known from Cambrian rocks. (Pleurotomaria
Sowerby, 1821, is not among them and indeed did not appear in the
Paleozoic.) These four are Sinuopea Ulrich, 1911 (pl. 2, fig. 1),
Schizopea Butts, 1926 (pl. 2, fig. 2) (=Rhachopea Ulrich and Bridge,
1931, and Roubidouxia Butts, 1926), and Dirhachopea and Taenio-
spira, both of Ulrich and Bridge, 1931. The anal emargination is a
deep, rounded sinus in Sinuopea and a deep angular sinus in Schizopea
and Dirhachopea, perhaps culminating in a short notchlike slit in the
latter. In Taeniospira there is a moderately deep slit and a typical slit
band. All four genera are known from beds no older than the latest
Cambrian Trempealeauan stage.

Six typical and unquestionable bellerophont genera are now known
from the Cambrian and will the neontologist please note that Bellero-
phon Montfort, 1808, is not among them. These are Owenella Ulrich
and Scofield, 1897, and Cloudia, Anconochilus, Sinuella (pl. 1, fig.
10), Sirepsodiscus (pl. 1, fig. 8), and Chalarostrepsis (pl. 1, fig. 12)
(all of Knight, 1947 and 1948). The first four have rounded sinuses
as anal emarginations, the fifth a deep V-shaped sinus, and the last
a deep slit. All these are of late Cambrian age. The earliest is
Strepsodiscus of the early late Cambrian Dresbachian stage, and
three of them, Strepsodiscus, Sinuella, and Anconochilus, occur earlier
than any known pleurotomarian genera. Also there are two isotro-
phically coiled genera, Coreospira Saito, 1936 (pl. 1, fig. 7), and
Cycloholcus Knight, 1947, both referred to the Coreospiridae. Al-
though neither has an anal emargination, the Coreospiridae are here
regarded as primitive bellerophonts. Corcospira first appeared close
to the boundary between the Lower and Middle Cambrian, probably
on the upper side.

There is also still another genus appearing still earlier and ranging
throughout the Cambrian that must be considered in this connection.
It is Oelandia Westergard, 1936, which is here placed in the Coreo-
spiridae. It will be considered more in detail on a later page.

In addition to the bellerophont genera discussed, three genera of
macluritoid gastropods occur in the last stage of the Upper Cam-

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

brian, the Trempealeauan, Scaevogyra Whitfield, 1878 (pl. 2, fig. 7),
Matherella Walcott, 1912 (pl. 2, fig. 10), and Kobayashiella Endo,
1937. All other known Cambrian gastropods are referable to iso-
pleuran monoplacophoran genera. Of these Helctonella (pl. 1, fig. 2),
and Scenella Billings, 1872 (pl. 1, fig. 1), both put in their appearance
along with Oelandia (pl. 1, fig. 5) in the Lower Cambrian and are
thus among the earliest gastropods known. In addition to these there
is that very puzzling, problematical group of gastropodlike shells,
Pelagiella Matthew, 1895, and its allies, that range throughout the
Cambrian. These, for reasons given later in this paper, may be re-
garded as an independent branch from some unknown gastropod
ancestor or they may not be gastropods at all.

Although not yet described or announced in the literature chitons
(Polyplacophora) are known from Upper Cambrian beds of the
Trempealeauan stage.

CLIMBING DOWN THE FAMILY TREE

THE PLEUROTOMARIAN-BELLEROPHONT BRANCH TO THE ISOPLEURAN
MONOPLACOPHORA

Continuing to proceed from the better known to the less well
known, we will work backward from the living pleurotomarians, from
which can be gleaned the basic anatomical details of the group, search-
ing step by step for fossil forms that may be taken for representatives
of the various stages in their evolution from their most primitive an-
cestral stock. The living pleurotomarians are referred currently to
the Pleurotomariidae, the Scissurellidae, the Haliotidae, and the
Fissurellidae.

In starting on our exploration it seems safe to assume that the
basic organization of the most ancient pleurotomarian was essentially
the same as that of its living representatives. Such a procedure per-
mits us to drop rapidly down the gastropod family tree or backward in
time something over 400,000,000 years to the late Cambrian when the
first known pleurotomarians lived, continuing all the while along a
branch that is easily recognized because its members show asym-
metrical coiling and because of the anal emargination, a slit, sinus, or
notch in the outer lip of the shell. From this vantage point in the
remote past we may examine our surroundings, particularly those a
little more ancient. The objects of our search are forms that resemble
the pleurotomarians very closely but are still more primitive.

Contemporaneous with the earliest known pleurotomarians and in

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 25

part preceding them are the bellerophonts.® Al! but the most primi-
tive are so very similar to the pleurotomarians in a number of signifi-
cant particulars that on comparative anatomy alone they must be
regarded as quite closely related. The shells of the bellerophonts are
coiled typically in a close spiral but the coiling is isostrophic rather
than helicoidal; the whorl cavity is, of course, very deep and the
two symmetrical retractor muscles are inserted one on each side deeply
within the aperture at the two ends of the columella in such a posi-
tion that their retraction would withdraw the head and foot within
the aperture ; there is an anal emargination, a U-shaped or V-shaped
sinus or a slit, just as in the contemporary pleurotomarians. In fact
the only obvious particular in which the bellerophonts differ from
pleurotomarians is that the coiling is isostrophic and the shell is a sym-
metrical spiral. Clearly then, the bellerophont, like the pleurotomarian,
was a prosobranch, but a symmetrical prosobranch. Since lateral
symmetry is a primitive character in the mollusks this is precisely
what one might expect in the immediate ancestor of the pleuroto-
marians which themselves retain more or less symmetrically paired
organs. It is commonly believed by neontologists that asymmetry is
an immediate and necessary result of torsion. No doubt the belief is
well founded in the sense that torsion precedes asymmetry and is a
prerequisite for it, but if the bellerophonts are prosobranchs as their
morphology strongly suggests and if torsion is the factor that dis-
tinguishes a bellerophont from an immediate laterally symmetrical
isopleuran ancestor, then, as the time factor insists, it is not necessary
to suppose that asymmetry was an immediate consequence. Of course
torsion furnished the unstable condition that ultimately led to asym-
metry.

Again surveying our surroundings, this time from the apparent base
of the bellerophont stem, we meet with two more genera that have
the characters one would expect of the very primitive bellerophonts.
One is Cycloholcus from the base of the Upper Cambrian Dresbachian
stage and the other is Coreospira (pl. 1, fig. 7) (both referred to
previously) from close to the boundary of the Middle and the Lower
Cambrian, probably on the upper side of the boundary. Both of these
forms are isostrophically coiled and thus in this respect are in accord

9 Some views expressed by Thiele, 1935 (p. 1125), and Wenz, 1938 (pp. 58-60),
on the probable anatomy and physiology of the bellerophonts will not, I think,
bear close scrutiny. Since I do not wish to interrupt the present argument to
give the reasons for my contrary views that the bellerophonts are prosobranchs
instead of primitively orthoneurous “Amphigastropoda,” as Thiele and Wenz
supposed, I am discussing the matter in an appendix to this paper.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

with the bellerophonts. Unlike previously recognized bellerophonts
there is no emargination in the lip that corresponds to the anterior lip.
This appears puzzling unless we remember that there is a feature
we were to look for in the primitive bellerophont.

Further exploration turns up the genus Oelandia (pl. 1, fig. 5),
a genus that may be interpreted most plausibly as being closely re-
lated to Cycloholcus and especially Coreospira (pl. I, fig. 7). Oelandia
has been associated commonly with Helctonella Grabau and Shimer,
1909 (pl. 1, fig. 2). For example, Wenz in 1938 (p. 88) places it in
the subfamily Helcionellinae in the family Tryblidiidae. There is
indeed a resemblance—a resemblance that appears to me to be hon-
estly come by but still not decisive taxonomically. In Helcionella the
apertural margins are in a flat or nearly flat plane. In Oelandia how-
ever the margins tend to be curved and one end, the end toward
which the apex bends, is considerably extended and often tilted up
as though to form a trainlike hood. If one attempts to think in terms
of soft anatomy this hood seems anomalous over the head but fits
nicely as a hood over the posterior train of the foot. Hence the ex-
tended or up-tilted end is here regarded as posterior. If this hypothe-
sis is accepted the apex is posterior and Oelandia may be considered
to be a very primitive isostrophic prosobranch gastropod in the Cor-
eospiridae, one that has not yet advanced to the stage of close coiling.
Of course the anal emargination has not yet appeared. Helcionella
remains in the Isopleura with the nontorted Monoplacophora. Oelandia
is an anisopleuran that may have been derived directly from Hel-
cionella and retains its characteristic ornamentation. Torsion may
have first taken place between these two genera in earliest Cambrian
or in pre-Cambrian time. This possibility will be discussed again.

The Coreospiridae are bellerophonts in respect to the shell coiled
or nearly coiled with lateral symmetry in a plane. In some other
respects they resemble more closely the group that we next meet
with, for although we have reached in Oelandia (pl. 1, fig. 5) close
to the beginning of the fossil record we have not fully surveyed its
contemporaries. There are still three kinds of gastropods or sup-
posed gastropods represented with Oelandia in the Lower Cambrian
rocks. One of these three, Pelagiella and its allies, seems anomalous
from any viewpoint and will be reserved for later discussion. The
other two fit into our picture very nicely. Both are cup-shaped and
show complete bilateral symmetry. Their ornamentation consists of
transverse undulations somewhat similar to those of Coreospira (pl. 1,
fig. 7) and Oelandia. One, the genus Scenella Billings, 1872 (pl. 1,
fig. 1), is cup-shaped with a conical shell and the apex tipped toward

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 27

the narrower end. The shell of the other genus, Helcionella (pl. 1,
fig. 2), is also cup-shaped, and includes species that are low and
broad as well as others that are high and narrow. In both the apex
points toward the narrower end of the aperture and in the high and
narrow species it is almost hooked. None of these have the hoodlike
train of Coreospira, Oelandia, and narrow bellerophonts in general.

Although we know nothing of the internal organization of either
Helcionella (pl. 1, fig. 2) or Scenella (pl. 1, fig. 1) by direct ob-
servation, their external features such as shape and ornamentation
suggest rather strongly that they belong to a family that continues
into the Devonian. Specimens of an Ordovician genus of this
family, Archaeophiala*® Perner (pl. 1, fig. 3), preserve the muscle
scars beautifully. The scars are strongly pigmented and for that
reason are unusually sharp and clear. (See Knight, 1941, pl. 3, figs.
3a-b.) These scars are 12 in number and are arranged in a ring deep
within the margin of the shell. Two of the scars are larger than the
others and are made up of three parts. These tripartite scars, which
occur at one end, may be regarded as compound and perhaps as rep-
resenting the scars of three muscles each. The other 10 scars are
simple and probably are the scars of single muscles. These 12 (or 16)
scars are in bilaterally symmetrical pairs. The pair of large compound
scars lies at the end toward which the apex lies and very nearly closes
the circle at that end. The scars of the other five pairs follow sym-
metrically on either side until the circle is nearly closed at the other
end. There is a line of much fainter, unpigmented scars outside of
the principle ring. The six (or eight) pairs of pigmented scars were
probably points of attachment for symmetrically paired muscles con-
necting the shell to the foot. One can hardly guess what function was
served by the muscles that made the more obscure scars outside those
of the main circle but these shadow scars appear to be characteristic
of the group.

Two exceedingly important inferences are suggested by the scars
of Archaeophiala (pl. 1, fig. 3). The first inference is that the soft
anatomy was bilaterally symmetrical throughout, that is to say the
animal had not undergone torsion. This is an inference primarily
from the complete bilateral symmetry of the paired muscle scars,

10] am employing Archaeophiala rather than Tryblidium to typify the gastro-
pods with paired muscle scars for the reason that its shape, which is essentially
that of Scenella and the lower, cup-shaped Helcionella, suggests that it is the
more primitive. Although their muscle scars are virtually identical, I am plac-
ing each in a separate family, as will be seen, since each seems to be a mem-
ber of a different series, each with its characteristic shape.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

supported by the lack of an area between scars at either end for a
pallial cavity. The second inference is that the end that has the large
compound muscle scars and toward which the apex lies is anterior.
This follows as probable from a corollary to the principal of cephali-
zation to the effect that ‘““heteronomous segmentation is an expression
of cephalization.” If one takes these two inferences together with
the previous inference that such Cambrian genera as Scenella (pl. 1,
fig. 1) and at least the lower, cup-shaped species of Helcionella (pl. 1,
fig. 2) are organized in a similar way we have a working hypothesis
as to the organization of these very important early forms. It seems
quite certain that the superficial resemblances of these Cambrian cup-
shaped forms to the living prosobranch patellaceans or capulids or
to the equally prosobranch Paleozoic platyceratids is as surely a mat-
ter of convergence as is the equally superficial resemblance of all of
them to the pulmonate ancylids.

It seems probable that a prerequisite for torsion was a reduction
in the hypothetical six or eight paired shell muscles to a single pair.
What better mechanism to give mutations accomplishing such a re-
duction survival value could have been devised than the development
of high, narrow shells, such as actually occurred in some Lower and
Middle Cambrian species currently referred to Helcionella (pl. 1,
fig. 4). In these the hypothetical six or eight pairs of muscles, if
present, would be crowded together. Perhaps, owing to this crowd-
ing, mutations that would effect the reduction of the six or eight pairs
to a single pair through the elimination of all but one of the pairs
would have survival value. If the suggested reduction actually took
place the foundation was laid for torsion. All that would be required
further is that through a genotypically small mutation the rudiments
of one muscle of the pair (the left one) should develop in the early
veliger larva earlier than those of the other. As has been shown by
Crofts (1937), the retraction of such a single asymmetrical “velum
retractor muscle” in the early veliger is what actually initiates torsion
in Haliotis. Undoubtedly when torsion first appeared in the remote
ancestors of Haliotis the same mechanism was responsible for it.

Both Helcionella (pl. 1, figs. 2 and 4) and Scenella (pl. 1, fig. 1)
appear in Lower Cambrian rocks. It seems probable that Helcionella
and Scenella had a common ancestor in early Cambrian or in pre-
Cambrian time. Chuaria Walcott, from pre-Cambrian rocks of the
Grand Canyon region, has been suggested as the most primitive an-
cestral gastropod but the only known specimens of the only known
“species,” all of which I have examined, are so very poorly preserved
that it is utterly impossible for me to recognize them as gastropods or

NO. 13 PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 29

anything else. The most I can say of the specimens is that they may
be organic in origin.

In descending the family tree we have passed from the earliest
forms that can be assigned to the Anisopleura with assurance, the
bellerophont cyrtolitids and sinuitids, such as Strepsodiscus (pl. 1,
fig. 8) and Sinuella (pl. 1, fig. 10) of the lower and middle Upper
Cambrian, through the probably anisopleuran Coreospiridae, to the
isopleuran Helcionella (pl. 1, fig. 2) and Scenella (pl. 1, fig. 1) of the
Lower and Middle Cambrian. In doing so we have passed along two
exclusively Cambrian limbs, the Coreospiridae and the Helcionellidae.
The Coreospiridae resemble the bellerophonts externally except that
there is no feature that can be assigned the function of the bellero-
phont anal emargination, The Helcionellidae resemble the Coreo-
spiridae except that the direction toward which the apex bends is
interpreted as anterior. As stated above, there are with the genus
Helcionella (pl. 1, figs. 2 and 4), as currently understood, species that
have a high shell with a strongly curved apex and others, more simi-
lar to the type species, with a low shell with the apex so short and
blunt that in some specimens it is almost an overstatement to say that
it is curved at all. These appear to make a continuous series. Our
hypothesis requires that torsion was initiated somewhere between
the untorted helcionellids and torted bellerophonts. The evidence for
one point in the chain as against another is not very compelling. I
have placed the dividing line between Helcionella (pl. 1, fig. 2) and
Oelandia (pl. 1, fig. 5), placing the former in the Isopleura with the
Monoplacophora and the latter in the Anisopleura with the bellero-
phonts. If anyone prefers to class the Coreospiridae with the Mono-
placophora or Helcionella with the bellerophonts, I cannot quarrel
too vigorously with the preference. There is insufficient evidence. As
the muscle scars, which might give more objective evidence, are un-
known in Helcionella and in both Oelandia (pl. 1, fig. 6) and Coreo-
spira (pl. 1, fig. 7) we are left with little but interpretations from
weak morphological data as basis for a decision, however tentative.
What little objective evidence there is lies in the similarity of the
ornamentation in the Helcionellidae and the Coreospiridae and in
differences in the apertural margins. This suggests that both of them
are allied to each other and to the Scenellidae where the ornamenta-
tion follows a similar pattern, but that for some reason, assumed to be
torsion, the apertural margins are different. At whatever point tor-
sion was introduced, our hypothesis requires that it was in the more
or less advanced descendants of Scenella and Helcionella that con-
ceivably retained a similar type of ornamentation.

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Just as the neontologists have employed restorations of the hypo-
thetical primitive mollusk with fruitful results, so the paleontologist
with even more actual data, the fossil shells, may employ them also.
Not only does the paleontologist have fossil shells that tend to sup-

Ventricle- - f=
Auricle— 7 4

Ctenidium---

Anus---43

FIGURE 7

a, Schematic restoration of Coreospira as a monoplacophoran isopleuran.

b, Schematic restoration of Coreospira as an isostrophic anisopleuran, a bel-
lerophont without an anal emargination. The latter seems a much more
plausible restoration than the former. Of course, neither restoration may
approximate the truth, but in that case Coreospira would probably not
have been a gastropod. It is understood that such organs as ctenidia,
auricles, etc., are paired in both restorations. The probable retractor
muscles are not shown.

port the scientific speculations of the neontologist but he has others
to which he may attempt to fit the soft parts of a generalized gastro-
pod and form judgments from the plausibility of the results as to
what the animal as a whole may have been like. Some of these hypo-
thetically restored gastropods. tend to fill gaps between the untorted

NO ES PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 31

monoplacophoran and the torted bellerophont which in turn connects
closely with the pleurotomarians.

For example, text figure 7 shows two restorations based on the
known shells of Coreospira (pl. 1, fig. 7). Figure 7, a, shows the shell
and hypothetical soft parts restored as an isopleuran monoplacophoran.
Figure 7, b, shows the same restored as an anisopleuran bellerophont.
Obviously the second yields a plausible picture of the probable re-
lationship of shell and soft parts. It looks comfortable. The mono-
placophoran restoration is too fantastic for even tentative acceptance.
Even though one should restore the soft parts to display more primi-
tive isopleuran features, a row of muscles, a very shallow posterior

FIcurE 8

Oelandia restored as a bellerophont even more primitive than Coreospira. It
presents a harmonious and plausible picture. A restoration as a mono-
placophoran is quite as unacceptable as is the same restoration of Coreospira.
As I have pointed out previously the trainlike hood over the posterior part
of the foot is a critical feature—a feature that is shared with several
bellerophont genera with narrow shells.

pallial cavity, etc., we still would have the coiled shell suspended
above the head in a most unacceptable fashion, as well as a narrow,
coiled visceral mass entirely incongruous on a monoplacophoran.
Surely it is difficult to accept Coreospira as other than a primitive
bellerophont.

Figure 8 shows a restoration of Oelandia (pl. 1, fig. 5), believed
to be a bellerophont even closer to the Monoplacophora than Coreo-
spira (pl. 1, fig. 7).

THE POLYPLACOPHORAN BRANCH

The procedure of working backward may be likened to selecting
one terminal twig of a tree from among very many, a twig on a

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

branch that by preliminary inspection took its origin far down on the
trunk, and then following that branch still farther down until one
is led by the process to what appear to be the roots. But our meta-
phorical tree, from preliminary inspection in very poor light (for let
us assume that we are feeling our way in the dark of the moon),
seems to have more than one main branch. One of these which we
will call the Polyplacophora, appears near the roots to lie close
to the branch that we have been tracing backward with apparent
success. Let us examine it further.

The chitons or Polyplacophora, far from abundant today, have al-
ways been rare in the fossil record. Nevertheless they are reported in
the literature as distributed throughout geological time from rocks
as early as Lower Ordovician and in the collections of the United
States Geological Survey housed in the United States National Mu-
seum are specimens of unquestionable polyplacophoran plates from
the late Upper Cambrian Eminence dolomite of Missouri. These
specimens, belonging to species and perhaps genera yet undescribed,
are nevertheless typically polyplacophoran in every detail including
the peculiar surface sculpture, common to all chitons, possibly to be
associated with the remarkable shell eyes, or aesthetes, developed in
this group.

Thus our leap backward in time along the polyplacophoran limb of
our metaphorical tree carries us almost exactly as far as our leap
along the pleurotomarian limb, to latest Cambrian time. However, we
find no obvious intermediate connections with any monoplacophoran.
Our only clue appears to be offered by the paired multiple dorsal
muscle scars of Archaeophiala (pl. 1, fig. 3) attributable by analogy
to Helcionella (pl. 1, fig. 2) and Scenella (pl. 1, fig. 1), possibly rein-
forced by what appear to be tubules in Tryblidium very similar to those
which carry the nerves for the aesthetes in chitons. In Archaeophiala
(and in Tryblidium) the number of pairs is six? but the pair of
large scars at the end regarded as anterior are compound and made
up of three smaller elements so that the basic number of pairs might
be regarded as eight. One might infer that the eight-segmented shell
of the polyplacophoran was merely the single shell of the mono-
placophoran separated into eight segments to correspond with the
eight pairs of shell muscles.

11Tt may be significant that the embryos of living polyplacophorans first
develop six shell plates. The other two, the terminal plates, are added at a later
stage (Garstang, 1920, p. 78).

NO. 13 PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 33

RECLIMBING THE TREE

For recapitulation it may be well to reverse our course and sum-
marize our results by ascending the pleurotomarian branch of the
family tree beginning with the Monoplacophora. We will still hold
to this one line, lest we go entirely astray, and we will arrive at the
present-day level along a limb with nothing more advanced than the
highly specialized relics of the once great pleurotomarian stock.

Throughout rocks of Cambrian age we find what appear to be
primitive gastropods with low, cuplike shells. The apex is subcentral
or anterior and there is no posterior train. All have rather coarse
transverse plicae or costae and finer ornamentation as well. They
are believed to have six (or eight?) symmetrical pairs of adductor
muscles and not to have undergone torsion. Typical of these early
Cambrian genera are Scenella (pl. 1, fig. 1) and those species of
Helcionella (pl. 1, fig. 2) that have the low cuplike form of the
genotype. Probably these or similar forms were in existence in late
pre-Cambrian time.

Concurrent with the more typical species of Helcionella (pl. 1,
fig. 2) are other species, that should probably be referred to another
as yet unnamed genus, which have very high, narrow shells (pl. 1,
fig. 4). It is possible that in these forms the adductor muscles were
so crowded that their number was reduced to a single pair, seemingly
a prerequisite for the initiation of torsion. Likewise in the early two-
thirds of Cambrian time are found species of Oelandia (pl. 1, fig. 6),
much like Helcionella externally but with an extended or up-tilted
margin under the apex that has the same shape as the posterior train
found in narrow bellerophonts. Accepting it as homologous, we then
must accept Oelandia (pl. 1, fig. 5) as having undergone torsion but
in most other respects to have retained at least some of the external
features of Helcionella. It is possible that it was the first bellerophont
and first prosobranch. Its apex is posterior but still not truly coiled.
However, close coiling is found in Coreospira (pl. 1, fig. 7) partly
contemporaneous with all these but appearing first a little later and
still without the anal emargination.

In late Cambrian time we find a number of bellerophonts each pro-
vided with an anal emargination: Strepsodiscus (pl. 1, fig. 8) and
Cloudia in the Cyrtolitidae; Sinuella (pl. 1, fig. 10), Owenella, and
Anconochilus in the Sinuitidae; and Chalarostrepsis (pl. 1, fig. 12)
in the Bellerophontidae. With this beginning the bellerophonts de-
ploy throughout Paleozoic time and have their last representatives
in the Triassic.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

In the latest Cambrian the first pleurotomarians put in their appear-
ance, mostly primitive pleurotomarians with either rounded or angular
sinuses, Sinuopea (pl. 2, fig. 1), Schigopea (pl. 2, fig. 2), and Di-
rhachopea. The anal emargination in Taeniospira is a true slit. The
anal emargination became a true slit in Early Ordovician time
in a number of genera, some as yet undescribed. During the re-
mainder of the Paleozoic the pleurotomarians proliferate greatly
and seemingly gave direct rise independently to a number of non-
pleurotomarian aspidobranch stocks and through these to most if not
all of the more advanced gastropods. They continue in declining num-
bers through the Mesozoic and Cenozoic and survive today in greatly
reduced numbers as relic families adapted to special environments.

In late Cambrian rocks in beds almost contemporaneous with those
containing the earliest known pleurotomarians, the first known poly-
placophorans appear, typical chitons in all respects. These also con-
tinue to the present day but always as relatively few forms mostly
adapted to rock clinging. They, like the anisopleuran branch, seem
to have been derived from primitive, untorted monoplacophorans but
through an entirely different set of modifications. The primitive iso-
pleuran condition continued, for in the polyplacophorans there was no
torsion, but the primitive single cuplike shell is replaced by eight
transverse plates. Perhaps these eight plates represent the primitive
shell which may have become divided transversely in accordance
with the possibly eight pairs of shell muscles.

EXPLORATION OF OTHER EARLY BRANCHES

In our climb down two branches of the gastropod family tree, arriv-
ing along both at the same main stem, we have followed what appears
to be a logical and straight course, paying no attention to other
nearby branches. But there are other nearby branches not too far
above the roots and it would be improper to leave them out of con-
sideration altogether, especially as the light is very poor.

THE PATELLACEA

First there is the branch that we will call the Patellacea. It is well
represented in our living faunas and goes far back into geological
time. The patellaceans include simple, cuplike shells that show ex-
ternally full bilateral symmetry and resemble very closely those that
we are here regarding as monoplacophoran isopleurans. However,
the anatomy and ontogeny of living representatives show unequivo-
cably that the symmetry of the shell is superficial and secondarily

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 35

derived. They are classified by neontologists as Archaeogastropoda in
the Prosobranchia. Although primitive in many respects, they show
in their soft parts and in their ontogeny both torsional and lateral
asymmetry. Can it be that in following our branch backward in time
we have become confused in the darkness and, instead of passing
from the earliest bellerophonts to monoplacophorans, we have stepped
across onto another superficially very similar but different branch?
Can it be then that what we are calling monoplacophoran isopleurans
are in truth nothing more than very ancient patellaceans? Except
for Wenz, most previous authors have so regarded them. However,
I think not, for there are characters in both groups, very obscure ones
to be sure, that seem to indicate the contrary.

The significant clues have to do with the scars of the shell muscles
in each group. In the patellaceans the muscle scars form a continuous
horseshoelike crescent, open anteriorly, for the shell muscle does not
intrude upon the region occupied by the anterior pallial cavity. The
shell muscle is composed of closely applied bundles of muscle fibers
and in some species this is reflected in the scar by knots, so to speak,
in the continuous scar that suggest the discrete scars of the typical
monoplacophorans. But these knots in the patellacean scar are not
symmetrically paired while the discrete scars of the monoplacophorans
are. They reflect the basic asymmetry of the patellaceans. Likewise
the anterior opening of the patellacean scar seems to reflect the torsion
of the primitively posterior pallial complex and cavity to an anterior
position above the head. Although there is in the patellaceans a very
thin scarlike line connecting the open ends of the horseshoe, it is
apparently not the scar of the pedal muscle but merely the line of
attachment of the mantle to the shell, analogous to the pallial line
of the pelecypods. The monoplacophorans are here conceived to have
included also forms with a continuous muscle scar, such as Archina-
cella Ulrich and Scofield, 1897, as well as those with discrete paired
scars, but in both types the scars have elements that close or nearly
close the circlet anteriorly and these elements seem to be continua-
tions of the scars themselves. This suggests that these forms, like the
Polyplacophora, do not have an anterior pallial cavity and supports
our inference that the Monoplacophora have not undergone torsion.
That the scars are narrower anteriorly may be accounted for if we
imagine that the muscles attached at this part are extensions from
the pedal muscles at each side arching over the head.

Although Wenz did not recognize them as such, it is my opinion
that the late Paleozoic genera, Metoptoma Phillips, 1836, and Lepe-
topsis Whitfield, 1882, are not monoplacophorans but are referable to

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

the Patellacea. Both have continuous, horseshoe-shaped muscle scars,
completely open at the anterior end. I would also assign Palaeo-
scurria Perner, 1903, to the Patellacea. Perner described and figured
for this genus an open horseshoe of almost discrete muscle scars but
I have examined the types of his genotype species and can find no
objective evidence for the existence of such a feature (Knight, 1941,
p. 231). Nevertheless, there is no direct evidence for any other sort
of scar, possibly because the matrix is too coarse to record such deli-
cate features. However, the shape of the shell is so similar to that
of Lepetopsis that I shall provisionally associate the two. The fossil
record of the Patellacea is then continuous from at least Mississippian
and perhaps from Silurian time to the present. I know of no Patel-
lacea from rocks earlier than Silurian, nor do I know any forms
transitional from pleurotomarian to patellacean unless the very im-
perfectly known Halophiala Koken, 1925, from Ordovician rocks
may be so regarded.

MACLURITES AND ITS ALLIES

Beginning in the early Trempealeauan stage of the Upper Cam-
brian and ranging into the Middle Devonian are a series of genera
that give the appearance, at least, of being coiled sinistrally. These
genera are here united taxonomically not only by the apparent sinis-
tral coiling, but by another feature as well. This feature, a difficult
one to describe, consists in most of these genera of a peculiarity of
the region surrounding the umbilicus or that part of the shell usually
called the base whereby the “basal” part of the whorl profile is rather
sharply arched, most conspicuously so where there is an open um-
bilicus. This sharp arching of the supposed basal part of the whorl
resembles a notch keel with an internal channel. In many forms it
is clearly the locus of a sinus in the lip. The following 17 genera,
most of them commonly regarded as sinistral, make up the group I
have in mind:

From the Upper Cambrian rocks

Kobayashiella Endo, 1937.
Matherella Walcott, 1912 (pl. 2, fig. 10).
Scaevogyra Whitfield, 1878 (pl. 2, fig. 7).

From Ordovician rocks

Antispira Perner, 1903.

Barnesella Bridge and Cloud, 1947 (p. 545).
Clisospira Billings, 1865.

Helicotis Koken, 1925.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 37

Laeogyra Perner, 1903.

Lecanospira Ulrich, in Butts, 1926 (pl. 2, fig. 8).

Lesucurilla Koken, 1898.

Maclurites LeSueur, 1818 (pl. 2, fig. 12) (= Maclurina Ulrich and
Scofield, 1897).

Macluritella Kirk, 1927.

Matherellina Kobayashi, (1033) 1937.

Mimospira Koken, 1925.

Palliseria Wilson, 1924 (pl. 2, fig. 11) (=Mitrospira Kirk, 1930).

Versispira Perner, 1903.

From Silurian rocks

Onychochilus Lindstrom, 1884 (pl. 2, fig. 9) (= Palaeopupa Foerste,
1893).

From Devoman rocks

Sinistracirsa Cossman, 1908 (= Donaldia Perner, 1903, preoccupied,
and Boycottia Tomlin, 1931).

Omphalocirrus Ryckholt, 1860 (=Coelocentrus Zittel, 1882, Poly-
enaulus Ethridge, 1917, and Arctomphalus Tolmachoff, 1926).

Thus I have grouped together (with one or two superficially dex-
tral genera) all the Paleozoic genera commonly regarded as sinistral
except Antitrochus Whidborne, 1891, which I refer tentatively to
the Trochonematacea, Agnesia Koninck, 1883, and Hesperiella Holz-
apfel, 1889, both pleurotomarians and possibly congruent, and Cam-
bodgia Mansuy, 1914, a pseudomellaniid. Other typically dextral
genera are known to have a few sinistral species, as well.

Up to this point I have spoken of the gastropods of the group we
are considering as “apparently” sinistral, that is to say, when the
shell is oriented in the arbitrarily conventional position '* with the
spire upward (or the umbilicus downward), the aperture is below
and to the left rather than to the right as in the vast majority of
gastropods. In the truly sinistral gastropod all organs of the body
are reversed in position from that of the dextral gastropod beginning
ontogenetically with the early cleavages of the egg. The reversal
appears to be the result of a mutation that may occur in some indi-
viduals of normally dextral species, or that has become fixed in the
heritage of some species in genera that are otherwise dextral, or of
a few entire families.

Sinistrality is well known among living gastropods but relatively
it is very rare. Likewise it is known among fossil Gastropoda. Of

12 | employ the illogical conventional orientation preferred by English, German,
and American authors.

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS It

the genera dealt with above Antitrochus and Cambodgia are probably
sinistral. The pleurotomarians Agnesia and Hesperiella present a
still different picture that I hope to discuss at another time. But in
all cases sinistrality is a deviation from the basic plan and seemingly
occurs only as the result of mutations that may or may not become
fixed in the heritage of a group. It occurs sporadically in various
only remotely related groups and is probably of no selective value,
positive or negative, to its possessor. The rarity of sinistral gastro-
pods is related to the primitive torsion and asymmetry of the Aniso-
pleura. Presumably it was of such a nature as to produce dextral
forms, and deviations from the plan require a relatively rare muta-
tion in which all parts of the organism at all stages were reversed.
Therefore the occurrence of a relatively large number of apparently
sinistral forms classifiable into a relatively large number of genera
very early in the history of the class is startling and affords grounds
for suspicion that these forms are not truly sinistral.

Among living gastropods there is another phenomenon very much
rarer than sinistrality which gives rise to a shell that has the appear-
ance of being sinistral but the organs of the anatomy are not reversed
from the position in dextral Gastropoda. The entire animal, including
both soft parts and shell, is actually dextral in this case, and the
shell is ultradextral or hyperstrophic, not sinistral. In other words,
the normal spire has sunk inward, as it were, and may even be coiled
in such a way as to produce an umbilicus. The normal base may be
flat or protrude to resemble a spire in every respect. Hence a shell
is produced that appears to be sinistral although it is actually dex-
tral. The “spire” of such a shell is homologous with the base of what
may be called a normal dextral shell and its “base” is homologous
with the spire. (See fig. 2.)

Hyperstrophy is exceedingly rare among living gastropods, occur-
ring most frequently as a specialization only in the embryonic nucleus
of some opisthobranch gastropods and in the adult stage of a few end
members of various highly specialized groups such as pteropods,
Ampullariidae, and pulmonates. If it were to occur in adults of spe-
cies with unknown soft parts it would be difficult to distinguish from
sinistrality except on collateral evidence. One line of collateral evi-
dence is that supplied by the peculiar angulation on the “base” of
these early Paleozoic shells. If we regard these shells as hyper-
strophic, the angulation is no longer anomalous. It becomes the trace
of the dorsal anal emargination. There is another line of collateral
evidence that is exceedingly pertinent to at least one of the Paleozoic
genera that is included in the group we are discussing and it seems

NO. 13 PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 39

very strong evidence indeed. This evidence is furnished by the oper-
culum which is preserved in this genus because it is calcified.

The gastropod operculum is basically corneous (conchyolin) but in
some groups the corneous operculum is partially or wholly calcified
and in some forms this makes the operculum very massive. Only
where it is calcified is the operculum of fossil forms preserved. The
embryonic operculum is a minute disk that grows by incremental addi-
tions to a margin or margins. Where the increments are added mark-
edly to one side of the margin as against the other sides, growth may
be in a spiral and such a spiral operculum, as seen on the external
face of the operculum in dextral gastropods, always grows from the
nucleus in a counterclockwise direction. In sinistral gastropods it is
clockwise. Now, in one of the genera of Paleozoic gastropods of the
group we are considering, the operculum was thick and calcified,
consequently it is not only frequently preserved but in some speci-
mens it has been found in place in the aperture. The genus is Mac-
lurites LeSueur, 1818 (pl. 2, fig. 12). In Maclurites the operculum
is in the form of an open spiral and the direction of coiling, as seen
on its outer face, is counterclockwise. Hence, as pointed out by S. P.
Woodward as early as 1854 (p. 202), the shell of Maclurites is not
sinistral, as has often been supposed, but dextral and hyperstrophic.
We do not know the operculum in any other of the genera included
in the group under consideration. Nevertheless, as I have endeavored
to show, the group we are considering appears to be a natural unit
and we may therefore with reasonable assurance attribute to the
other genera the character of hyperstrophy that the angulation on
the “base” of the whorls suggests and that the operculum of Mac-
lurites seems to confirm.

Assuming that we have solved the problem of the coiling in
the group under consideration, namely that it is hyperstrophic dex-
tral rather than sinistral, we are faced in consequence with an even
more difficult problem. What is the meaning in terms of soft anat-
omy and of phylogeny of a rather large group of dextral hyper-
stropic forms introduced so very early in the history of the Gas-
tropoda? Since this group seemingly became extinct before the
close of Devonian time, it left no recognized descendants among liv-
ing gastropods that might throw some light on its organization. As
suggested above, hyperstrophy is very rare among living gastropods
and occurs only as a secondary acquisition in groups far removed
from any possible connection with our early Paleozoic group. We
can only surmise what the anatomy of the soft parts of the Paleozoic
forms might have been. The shells are coiled and coiled asymmet-

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

rically. These facts suggest torsion and possible asymmetry in_ the
primitively paired organs. But, as the shells are hyperstrophic and
appear in the fossil record shortly before the first known pleuroto-
marians, the asymmetry possibly may be very different from that of
the main line of gastropod descent. Figure 9g shows hypothetical
restorations of the hyperstrophic genus Palliseria (pl. 2, fig. 11) a
close relative of Maclurites (pl. 2, fig. 12). Accepting the notch keel
surrounding the umbilicus as the locus of the anus we find very little
room for a right ctenidium and tentatively assume that this and its
associated organs had been lost. The operculum of Maclurites is not
only that of a dextral shell but it shows a startling resemblance to
that of the recent Nerita in that there are points of attachment for a
pair of retractor muscles. A single pair of retractor muscles is a
primitive feature shared with the bellerophonts, the more primitive
pleurotomarians and the neritaceans. The line of speculation that
seems most plausible to me is that this group branched off from the
early bellerophonts at some such stage as is represented by Strepso-
discus (pl. 1, fig. 8), a bellerophont that commonly shows some asym-
metry in a sinistral or hyperstrophic sense. This would accord with
chronogenesis, for Strepsodiscus precedes in the fossil record Scaevo-
gyra (pl. 2, fig. 7), the earliest hyperstrophic gastropod, and both
precede the earliest pleurotomarian. If this is true the somewhat
angular ““base” of Scaevogyra’s whorl is homologous with the angular
dorsum of Strepsodiscus. It is further supposed that asymmetry
arose in the group under consideration as an early genetic response
to the mechanical difficulties of isostrophic coiling as in the main line
of gastropod descent that began with the pleurotomarians, but inde-
pendently and probably in a somewhat different way, and that this, in
turn, resulted in the asymmetrical, hyperstrophic shell.

PELAGIELLA AND ITS ALLIES

Doubts have been expressed that the members of this group are
actually gastropods in spite of the very close resemblance of their
shells to those of gastropods. Thus Wenz wrote in 1938 (p. 95) of
Pelagiella: “Systematische Stellung Fraglich; vermutlich uberhaupt
nicht zu den Gastropoden gehorig.” The late Dr. E. O. Ulrich is
said to have held the opinion that they were “pteropods,” that is to
say, allied to the hyolithids.t* The hyolithids are no longer regarded
as true pteropods or even as gastropods for that matter. The true
pteropods are highly specialized opisthobranch gastropods of Tertiary

13 Oral communication from Dr. Josiah Bridge.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 4I

FIGURE 9

Three views of a restoration of Palleseria longwelli (Kirk), about X 4.

a, View of the left side. Note the direction of coiling and the protruding,
spirelike base.

b, Anterior view.

c, View of the right side. Note here and on the anterior view (b) the
umbilicus occupying the side where a spire would be in a dextral ortho-
strophic gastropod. Note especially that the ridge surrounding the
umbilcus is the locus of a notch. This notch, an anomalous feature if
the shell is regarded as sinistral, is believed to be the anal emargination.
If this is correct there is very little space for the primitive right ctenidium
and probably it has been lost.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

and Recent times, in many respects the farthest removed from the
primitive stock of any of the class.

Matthew, in erecting the genus in 1895, seemed to regard Pelagiella
as a heteropod. The heteropods are again a highly specialized group
of late Cretaceous to Recent times, although being prosobranch, not
quite so far removed morphologically from the primitive stock as
the opisthobranch pteropods. These quite unacceptable assignments
serve to accentuate the difficulties in finding a place for these forms
in the Gastropoda. My own difficulties derive from the fact that if
Pelagiella and its allies are gastropods, the hypotheses I have been
setting up cannot include them, except peripherally. Otherwise these
hypotheses must be abandoned or extensively modified in such ways
that they would meet with greater difficulties in other directions than
the difficulty presented in removing this obviously questionable group
to a peripheral position or eliminating it from the Gastropoda alto-
gether.

Pelagiella and its allies occur in company with the first recorded
monoplacophoran gastropods in the rocks of early Cambrian time and
are not only coiled but asymmetrically coiled. Thus, if they are
gastropods, they appear superficially to have advanced well beyond
the isopleuran monoplacophoran stage, the anisopleuran isostrophic
stage, and even beyond the pleurotomarian stage of the main line of
gastropod evolution. Therefore, since they are contemporaneous
with the earliest known monoplacophorans, and the earliest and most
primitive known representatives of one of the two hypothetically
more primitive stages and precede the others by a considerable in-
terval of geological time, the difficulties are obvious.

I do not wish to consume space in laboring the problem at too great
length for, with our present lack of knowledge of their anatomy and
even of their conchology, it is not soluble. I am not prepared to
abandon the hypotheses as to the derivation of the main lines of
gastropod descent until other hypotheses are presented that better
explain the observed facts. Although it is recognized that the fossil
record is imperfect, I am not prepared to assume that it is so very
faulty that its bearing on the broader aspects of chronogenesis is to
be set aside. There are various other possibilities that might be called
upon to account for Pelagiella and its allies as gastropods. For ex-
ample, can it be that they represent a branch from the monoplaco-
phoran stock that acquired torsion and asymmetry independently in
pre-Cambrian time, perhaps bypassing an isostrophic stage? There
is little evidence one way or another but I think it extremely unlikely.
Or can it be that they are monoplacophorans that carried their tend-

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 43

ency to coil anteriorly to such a point that the resulting inconven-
ience of a coil poised above the head gave survival value to any muta-
tion that set the coil asymmetrically to one side, as it were? This
again appears to be extremely unlikely. It would require a sym-
metrically coiled predecessor and no such form is known.

Finally then, I share the doubts of my predecessors, expressed or
implied, that Pelagiella and its allies are gastropods, but I shall go
further than they have gone in that I shall not attempt to force them
into a phylogenetic classification of the gastropods that appears to
have no place for them. I shall very tentatively assign them the
peripheral position of a shoot from the same pre-Cambrian root as
the main trunk of the gastropod family tree originating obscurely in
pre-Cambrian and of otherwise little-understood affinities. It would
be helpful if I could assign them elsewhere in the animal kingdom,
but I cannot do so.

Before leaving the subject of Pelagiella and its allies it may be
well to review briefly and in general terms their chief characteristics.
The shells are coiled and of from one-half to about three whorls. The
coil is always asymmetrical. For the most part they are small and
many are minute, a millimeter or two in diameter. The apical end of
the whorl, the nucleus, appears in some to be laterally flattened and
somewhat blunt, reminding one of the tip of a ram’s horn, and in
some forms slightly swollen. The whorls are ovoid in section, the
narrow end of the ovoid being at the periphery. The spire is always
low, varying in that respect from depressed to umboniform. In the
forms with a depressed spire the base is arched; in those with an
arched spire it is flattish. The shells of any one species appear to
be rather variable and it is probable that both dextral and sinistral
forms occur in some species. The ornamentation consists of fine,
faint lines of growth and, on some forms, a single faint revolving
lira, seemingly both above and below the periphery. The growth lines
are somewhat drawn back at the rounded periphery, thus suggesting
a broad, peripheral sinus. No operculum is known and there is no
information on muscle scars. In some specimens of Pelagiella,
Matthews reports and figures a groovelike constriction in the shell
(or its steinkern?) close to the apertural margin. It is not present on
Matthew’s primary types of Pelagiella atlantoides (Matthew), the
genotype (Knight, 1941, p. 237), but does occur on rare specimens
subsequently assigned to the species by Matthew. It may be a mark
of maturity or old age. Possibly the constriction is seen only on the
steinkern in which case it might mean only that the apertural margin

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

was thickened within and the thickening is invisible on the outer
surface of the shell.

This group, in which I include Cambrian species mistakenly referred
by authors to such genera as Straparollina Billings, Straparolus Mont-
fort, Euomphalus Sowerby, Raphistoma Hall, Ophileta Vanuxem,
and Platyceras Conrad, is in urgent need of intensive study, as are
all Cambrian gastropods and gastropodlike forms for that matter.
Several names have been proposed for supposed genera, mostly on
the basis of quite inadequate studies. Besides Pelagiella Matthew,
1895, there are Parapelagiella Kobayashi, 1939 (p. 287), Proto-
scaevogyra Kobayashi, 1939 (p. 286), and Proeccyliopterus Kobay-
ashi, 1939 (p. 286). The last three seem to be erected on characters
of very doubtful value or are differentiated from Pelagiella on mis-
taken.concepts of the characters of Pelagiella itself.‘* Still another
genus, Semicircularea Lochman, in Lochman and Duncan, 1944
(p. 44), was erected for the forms with only about one-half whorl
often misidentified as Platyceras by previous authors. Pelagiella and
its allies range throughout Cambrian and perhaps into early Ordovician
time.

TAXONOMIC CONCLUSIONS

As a result of our findings on our descent of the family tree and
of the paleontological and neontological considerations given above,
we have arrived at tentative hypotheses that force on our attention
certain taxonomic conclusions. The first is that, since the mono-
placophoran gastropods seemingly share with the polyplacophorans
the basic isopleuran plan of organization, the two should be brought
more closely together than has been customary in most classifications.
The second is that, since the Anisopleura as the result of mutation
arose stiddenly from a monoplacophoran ancestor, and since certain
anatomical features of both are very similar, the relationship between
them is too close to permit them to be arranged in separate classes
comparable in degree of differentiation to the other molluscan classes.
The third is that, although the isopleurans and the anisopleurans
should be placed in a single class, the gulf between them, both ana-
tomically and in time, is profound and that, therefore, it seems ap-
propriate to rank each as a subclass. On the basis of these three
considerations we present the following revised definitions of the class

14 Tt is unfortunate that the belief that the supposed characters of two of them
suggested that they were ancestral to later genera of quite different affinities and
led to the fixation of those ideas in the names given them. The proposal of
names embodying phylogenetic concepts is most unwise.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 45

Gastropoda and its major subdivisions, the subclasses Isopleura and
Anisopleura. There are also other conclusions as to the subdivisions
of the next lower rank but consideration of these is postponed until
the class and the two proposed subclasses are dealt with.

Class GASTROPODA Cuvier

So great is the range of special morphological modifications in the
class that it is exceedingly difficult, if not impossible, to draw up a
brief diagnosis that will cover all gastropods without excluding some
forms that clearly must be included. The return of the isopleurans
to the class, however necessary it appears, increases the difficulties,
for we thereby reduce the convenient criteria of torsion and of a
single shell to a status diagnostic of subdivisions of lower rank.

The gastropods may be defined as mollusks with a differentiated
head, a flat creeping foot, and a single basically conical shell. In a
few gastropods specialized for free swimming the foot may be modi-
fied into finlike organs, in the polyplacophorans the primitive single
shell has been divided transversely into eight segments, and in some
highly specialized forms the shell has disappeared in the adult. In
many others the cone is attenuated and coiled. Primitively marine,
they have become adapted also to fresh waters and to terrestrial life.
They are found at nearly all latitudes and nearly all altitudes from
the depths of the oceans to high mountains. They appear in the fossil
record in Lower Cambrian rocks and are flourishing today.

Subclass IsopLeura Lankester.—Gastropods that retain through-
out life both in the shell and in the soft anatomy the primitive bi-
lateral symmetry of the class. They are entirely marine and always
rare. They first appear in the fossil record in Lower Cambrian rocks
and carry through to the present. They probably originated in pre-
Cambrian time.

Subclass ANIsopLEURA Lankester.—Gastropods that undergo tor-
sion during the veliger stage. The Anisopleura are often abundant
and are tremendously diversified in morphology and in habitat. They
first appear in the fossil record as primitive forms in Lower Cam-
brian rocks and are flourishing today.

MAJOR DIVISIONS OF THE SUBCLASS ISOPLEURA

Order PotyptacopHora.—Isopleuran gastropods with the shell
made up of eight plates arranged along the midline of the dorsum;
head not provided with eyes; shell eyes, or aesthetes, may be present.
Polyplacophora range from late Cambrian time to the present, are
always marine and relatively rare.

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

The subdivisions of the Polyplacophora will not be considered here.

Order MonorL_acopHora.—lIsopleuran gastropods with a single
conical shell with the apex subcentral or pointed forward; some pos-
sibly with aesthetes. Marine, Lower Cambrian—Devonian,

Before considering the subdivisions of the Monoplacophora it may
be well to repeat that I do not consider Discinella Hall, 1871, Mober-
gella Hedstrém, 1923, or Barella Hedstrom, 1930, to be monoplaco-
phoran gastropods but hyolithoid opercula. Conchopeltis Walcott,
1879, I regard as probably a scyphozoan and certainly no mollusk.
Chuaria Walcott, 1899, is entirely problematical. (Knight, 1941,
p. 20.)

Family Trypiip1ipaE Pilsbry, 1899

Subfamily PALAEACMAEINAE Grabau and Shimer, 1909

Relatively low to high, cap-shaped shells with apex subcentral to
slightly anterior. Muscle scars (observed only in Archaeophiala)
discrete and arranged in six (or eight?) symmetrical pairs; orna-
mentation basically concentric undulations.

Genera

Scenella Billings, 1872 (pl. 1, fig. 1) (=Parmophorella Matthew,
1886), Cambrian.

Helcionella Grabau and Shimer, 1909 (pl. 1, fig. 2), throughout the
Cambrian.

Palaeacmaea Hall and Whitfield, 1872, Upper Cambrian.

Archacophiala Koken, in Perner, 1903(pl. 1, fig. 3) (=Scaphe Hedstrom,
1923, Scapha Hedstrom, 1923, Patelliscapha Tomlin, 1929, and
Paterella Hedstrom, 1930), Ordovician.

Calloconus Perner, 1903, Lower Devonian.

In the Silurian what appears to be a new genus hitherto unrecog-
nized is represented by Palaeacmaea? solarium Lindstrom, 1884

(p. 59).
Subfamily TRYBLIDIINAE Pilsbry, 1899

Spoon-shaped shells with the apex at or overhanging the anterior
end. Muscle scars (observed in Tryblidium, Pilina, Drahonura, Prop-
lina, and partially in Cyrtonella) essentially similar to those of the
foregoing family ; ornamentation concentric-lamellar or radiating.

Genera

Tryblidium Lindstrom, 1880, Silurian.
Cyrtonella Hall, 1879, Devonian.1®

15] have given my reasons for including Cyrtonella Hall in the Tryblidiidae
elsewhere (Knight, 1947b, p. 267).

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 47

Helcionopsis Ulrich and Scofield, 1897, Ordovician.

Drahomira Perner, 1903, Ordovician.16

Vallatotheca Foerste, 1914, Ordovician.

Pilina Koken, 1925, Silurian.

Proplina Ulrich and Bridge in Kobayashi, 1933, Upper Cambrian—
Lower Ordovician.17

Family HyPsELocoNnIDAE, new

Narrowly conical shells with the apex over the narrower (anterior ?)
end but tilted slightly backward. Ornamentation growth lines or
faint radiating undulations; muscle scars unknown.

Genera

Hypseloconus Berkey, 1898, Upper Cambrian-Lower Ordovician.18
Pollicina Holzapfel, 1895, Ordovician.

Family ARCHINACELLIDAE, new

Low conical shells with the apex at or overhanging the anterior
end. Ornamentation growth lines or radiating lirae; muscle scar a
broad, continuous ring, narrowing in front where it passes below the
apex. Ordovician.

Genera

Archinacella Ulrich and Scofield, 1897.
?Ptychopeltis Perner, 1903.!®

It is possible that Helcionopsis will find a place here rather than
with the Tryblidiidae when its muscle scars are discovered.

Order ApLAcopHoRA.—In this order there is no shell, and it is
consequently unknown as a fossil. I have no comments but retain it
here.

16 Drahomira is a name published, but not adopted, by Perner, 1903 (p. 23,
footnote) for Tryblidium glaseri Barrande in Perner, 1903 (p. 23), genotype
by monotypy. This name was overlooked by me in the preparation of “Paleozoic
Gastropod Genotypes” (Knight, 1941). Seemingly it is the valid name for a
distinct genus of this family.

17 The muscle scars of Proplina cornutaformis (Walcott), the genotype and
only species referred to the genus in published literature, are unknown. How-
ever, the material assembled for a monograph on Ozarkian and Canadian gas-
tropods by E. O. Ulrich and Josiah Bridge is available to me and several
species referred to the genus show them clearly.

18 For comments on the supposed multiple paired muscle scars of Hypselo-
conus see Knight, 1941 (p. 158).

19 Although Perner described a scar for Ptychopeltis, examination of his
specimens failed to disclose valid evidence for it (Knight, 1941, p. 288).

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

Susctass ANISOPLEURA LANKESTER

Except for reviving Lankester’s subclass Anisopleura, equivalent
without the Monoplacophora (Tryblidiacea of Wenz) to the class
Gastropoda of Wenz, 1938, I am now proposing few changes. To
the Bellerophontacea, which are retained as Prosobranchia, are added
the family Coreospiridae with the genera Coreospira Saito (pl. 1,
fig. 7), Cycloholcus Knight, and Oelandia Westergard (pl. 1, fig. 5),
but not without a residuum of doubt, and the superfamily Macluritacea
is erected in the Prosobranchia. The Macluritidae of Wenz form its
nucleus and other families composed of related elements are as-
sembled with it. The revised taxonomy of the group will be presented
as a part of another paper. The included genera are listed on pages
36-37 of this paper. In all other respects the Anisopleura are left as
Wenz had them but because of lack of opportunity for intensive study
rather than because of detailed endorsement of his arrangements,

APPENDIX
INTERPRETATION OF THE BELLEROPHONTS
AMPHIGASTROPODA VS. PROSOBRANCHIA

The genus “Bellerophon” of the older workers and some neon-
tologists (now expanded to a superfamily, the Bellerophontacea, with
four families and something like fifty genera and subgenera) has
been difficult to understand and to classify. Its isostrophic habit of
coiling is almost unique in the Gastropoda. This and the fact that
the entire superfamily has been extinct since Triassic time and affords
no living examples from which soft parts can be demonstrated have
seemingly left us with little information to go on. The broad mor-
phological pattern of the soft parts must be inferred since it cannot
be observed directly.

De Koninck in 1883 (p. 121) reaffirmed on a more rational basis
his suggestion of 1843 (p. 337) that the bellerophonts were proso-
branch gastropods. Before 1883 the bellerophonts had been regarded
as cephalopods, or as heteropod gastropods. Some specialized forms
such as Pterotheca, originally described as brachiopods or pelecypods,
have been regarded as pteropods. Since that time they have been
classified as prosobranch with the Docoglossa, or as a separate class
of Mollusca, the “Amphigastropoda.” I can subscribe to none of
these views except perhaps the main thesis of de Koninck in 1883,
although not to the details.

As stated previously, Wenz’s great contribution to theory in 1938

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 49

(p. 59) was the idea that the Tryblidiacea were primitive untorted
gastropods. This I applaud and accept. But he also regarded the
bellerophonts as untorted gastropods similar to the Tryblidiacea, on
the grounds of symmetry and an unsupported assumption that the
slit and band are not to be compared with the seemingly homologous
feature in the pleurotomarians. On this point I must part company
with him. Curiously, if I read the story aright, Wenz seems to be
following part way in the footsteps of many of his predecessors, who
regarded the bellerophonts as prosobranchs on the grounds of the
following chain of reasoning: The early cup-shaped shells are sym-
metrical and resemble the living patellids; therefore they are to be
classified with the latter as Docoglossa and prosobranchs. The bellero-
phonts are also symmetrical ; therefore they are closely related to the
patellids and are also Docoglossa and prosobranchs.

But Wenz in recognizing the early cup-shaped mollusks, the Try-
blidiacea, as nontorted gastropods changed the first premise of the
customary chain of reasoning and the bellerophonts, still linked with
these early cup-shaped shells, are, to Wenz, like them nontorted
gastropods.

The weakness in both lines of argument is the overvaluing of the
symmetry of bellerophonts as a criterion of relationship to the sym-
metrical cup-shaped shells whether patellids or tryblidians, the under-
valuing of the many manifest differences between the bellerophonts
and either of the other two, and the undervaluing of several manifest
anatomical homologies between the bellerophonts and the asymmetri-
cal but coiled prosobranch pleurotomarians.

Fundamental to the undervaluing of bellerophont-pleurotomarian
homologies is a failure on the part of Wenz and some neontologists
to recognize that torsion and the development of lateral asymmetry
are two distinct processes. Undoubtedly torsion set up unstable con-
ditions that favored the natural selection of mutations, such as lateral
asymmetry, that would result in a more efficient organism, but lateral
asymmetry is not to be confused with torsion or what I have called
torsional asymmetry. Although torsion is a prerequisite for asym-
metry, asymmetry does not necessarily follow from it. It is as though
this school of thought believes that the muscular pull that initiates
torsion also distorts the lateral symmetry. I know no evidence that
supports such a view. It is true, of course, that except in the Isopleura
all gastropods living today show lateral asymmetry at some ontoge-
netic stage and the lateral asymmetry is initiated in the veliger larva
immediately after torsion. But it does not follow that it was always
universally thus. The view that asymmetry was the immediate or

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

concommitant mechanical result of torsion and in consequence became
a part of gastropod heritage smacks somewhat of Lamarckianism and
in any case cannot be sustained.

Returning to Wenz’s views, in 1938 they seem to have been ap-
proximately as I have stated them above. On the basis of the lateral
symmetry alone he felt that the bellerophonts were closely related to
the tryblidians and consequently had not undergone torsion. Al-
though he gave no systematic expression to these views at that time,
classifying both the Tryblidiacea and Bellerophontacea as Proso-
branchia, it was his opinion that both were probably out of place in
that position.

Feeling insecure as to his interpretation of the bellerophonts he
was quite rightly searching for corroborative evidence, and he felt
that the discovery in the bellerophonts of multiple, paired dorsal
muscle scars like those of the tryblidians would be strong supporting
evidence, as indeed it would. In 1937 he wrote to me asking what I
knew of bellerophont muscle scars and that started the chain of events
about which I wrote ten years later (Knight, 1947). Briefly, a speci-
men of the supposed but somewhat atypical bellerophant Cyrtonella
mitella (Hall) was discovered which seemed to support fully Wenz’s
views in that the unmistakable record of two pairs of dorsal muscle
scars (not three as Wenz wrote) was clearly visible on that part of
the steinkern that was exposed. Possibly other scars may be covered
by matrix. Feeling that his views as to the close relationship of the
bellerophonts and tryblidians were fully vindicated, Wenz published
his paper giving systematic effect to those views by employing for
them a subclass, the Amphigastropoda (Wenz, 1940).”°

An interesting point about tryblidian muscle scars noted in Try-
blidium, Archaeophiala, and Cyrtonella is that each scar has on the
side toward the margin of the shell a smaller, fainter scar as though
it were the shadow of the scar cast before it. Wenz, who had never
seen the specimen of Cyrtonella mitella he figured, misinterpreted a
pair of these shadow scars, shown in the photograph sent him by Yang,
as a principal scar. This is why he mistakenly reported three visible
pairs of scars. The physiological significance of the “shadow scars” is

20 Actually the Amphigastropoda consisting of only the bellerophonts, was
erected as a new class of mollusks by Simroth in 1906 (p. 839), who was fol-
lowed by Thiele in 1935. Simroth’s course, and especially Thiele’s, was sup-
ported only by the gratuitous assumption that the soft anatomy was without
torsion and bilaterally symmetrical. Amazingly, Thiele assumed also “eine
anliche schwimmende Lebensweise—wie die Nautiliden” (Thiele, 1935, p. 1125)
in which he was followed by Wenz.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 51

obscure but their presence is an additional evidence that Cyrtonella is
a tryblidian.

Although the paired dorsal muscle scars on the specimen of
Cyrtonella were discovered in my laboratory and although it was on
my suggestion that Yang disclosed the discovery to Wenz, my views
as to its significance were quite different from those so promptly pub-
lished by Wenz in 1940. The more probable interpretation that
Cyrtonella, a genus that was even then somewhat doubtfully placed
in the Bellerophontacea and quite as easily interpreted as a tryblidian,
should be placed in the Tryblidacea instead of the Bellerophontacea
seems never to have occurred to Wenz.

Fortunately I was able to discover the muscle scars of two unques-
tionable bellerophont genera a few years later, Sinuwites (pl. 1, fig. 11)
and Bellerophon (pl. 1, fig. 13) (Knight, 1947). They consist of a
single symmetrical pair. Each muscle was attached to the opposite end
of the colummella about one-half whorl within the aperture, a posi-
tion that would permit them to serve effectively as pedal retractors.
They are not dorsal and not multiple pairs. Both of those facts are
seemingly fatal to Wenz’s arguments as to the closeness of the rela-
tionship between the tryblidians and the true bellerophonts.

Wenz displays a number of views to which I must take exception.
For example, he accepts the wholly conjectural and long-rejected
views of Lang (1891) as to the gradual development of torsion in
the gastropods. He treats the bellerophonts and pleurotomarians as
being present in early Cambrian rocks. In terms of genera recog-
nized by him, neither appeared until late Cambrian time. Under the
influence of his overestimate of the significance of external lateral
symmetry in the bellerophonts he fails to even consider the close
homologies between bellerophonts and pleurotomarians. Finally, he
seems to hold the view first proposed by Deshayes in 1830 (p. 135)
and abandoned by most students well before the close of the nine-
teenth century that the bellerophonts “tended toward a freely swim-
ming, nektonic mode of life” (translation from Wenz, 1938, p. 59).
I know of no evidence whatever that would support such a view and
would be interested indeed to learn of a molluscan swimming mecha-
nism that would be powerful enough to sustain the massive shells
of some bellerophonts above the sea bottoms. It seems highly prob-
able that the bellerophont foot conformed in general to the pattern
shown by other Archaeogastropoda. It was adapted to creeping, not
to swimming.

I regard the bellerophonts as prosobranch Archaeogastropoda close
to and probably ancestral to the pleurotomarians which they precede

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

in the fossil record. The bellerophonts share with the pleurotomarians
(1) a shell that typically has deeply hollow, usually closely coiled
whorls, (2) a sinus or slit which, if a slit, generates a slit band, (3) a
single pair of lateral retractor muscles,” and (4) seemingly a single
pair of each, of ctenidia, auricles, etc. They differ principally in that
the coil of the bellerophont shell is bilaterally symmetrical (iso-
strophic) and that of the pleurotomarian shell is an asymmetrical
orthostrophic helicoid, in my view a difference of little significance
for classification but of profound import for understanding gastropod
evolution.

Comparing the bellerophonts with the tryblidians we find they have
one feature, and only one, in common: externally the shell of each is
bilaterally symmetrical. But in respect to the first three categories
in the foregoing paragraph, the tryblidians have (with a very few
exceptionally high conical shells) (1) a shallow cup- or spoon-shaped
shell with the apex bent toward one end, but no coiling, (2) no sinus
or slit, and (3) multiple (usually six or eight) symmetrical pairs of
dorsal muscle scars.

Let us look for a moment at these points of agreement and dis-
agreement. The agreement between the bellerophonts and pleuro-
tomarians on points 1 and 3 can only mean that we have a shell with
a deep body cavity into which the head and foot can be withdrawn
by the retractor muscles which are properly placed in both for the
operation. Point 2, the sinus or slit can only be an anal emargina-
tion, a feature that is known otherwise only in prosobranchs and espe-
cially in the Archaeogastropoda, and which is accepted by many
neontologists such as Garstang, Yonge, Crofts, and many others as
an adaptation for sanitation after torsion had created a need for it. The
bilateral symmetry of the shell can no more be considered a charac-
ter of subclass or even ordinal rank than that same symmetry can be
employed to link two groups so different on other points as the
bellerophonts and tryblidians.

Need we continue to point further fundamental differences be-
tween the pleurotomarians and bellerophonts on one hand and the
tryblidians on the other, differences such as the impossibility of at
least the low, cuplike tryblidians pulling their head and foot into the
shell? In that respect they probably resembled the chitons and the
secondarily symmetrical fissurellids and patellaceans. Since the muscle

21 The living pleurotomariid genera or subgenera Perotrochus Fischer,
Entemnotrochus Fischer (pl. 2, fig. 4), and Mikadotrochus Lindholm, “Pleuro-
tomaria” of authors, have only a single retractor muscle, although other living
pleurotomarian genera have a pair.

NO. 13 PRIMITIVE FOSSIL GASTROPODS—-KNIGHT 53

scars of the high and narrow species of Helcionella (pl. 1, fig. 4)
in the tryblidians have never been observed, it is more difficult to
speculate profitably as to whether the muscles are so placed that they
could or could not have withdrawn the head and foot into the shell.
It is possible that they could, especially if, as I am suggesting, the
reduction of the pedal muscles from eight pairs to one pair may have
occurred first in them. Need it be pointed out to those who regard
the anal emargination in the bellerophonts as posterior that no known
or reasonably imaginable nontorted mollusk has or needs a slit or
sinus to provide egress for the contaminated water of a posterior
anus? Certainly the chitons and the tryblidians do not. Again, have
those who infer that bellerophonts are primitively orthoneurous non-
torted ““Amphigastropoda” ever tried to imagine the animal with its
large, heavy coil anterior and overhanging the head? The shell in
many bellerophont species is not only thick and heavy but may carry
a massive parietal callus as well. To me such an arrangement appears
highly improbable, bordering indeed on the fantastic. It is suggested
that the reader turn to figure 7, a, on page 30, where a restoration of
a smail very primitive bellerophont, with no parietal callus, is pre-
sented as though it were primitively orthoneurous and exogastric,
may help him to visualize it.

In summary, it appears to me that the evidence for the view that
the bellerophonts were prosobranchs, is very strong and the evidence
that they were primitively orthoneurous “Amphigastropoda” very
weak indeed.

Soft anatomy of the bellerophonts——We now know enough of bel-
lerophont shell morphology and enough of the morphology of living
examples of the obviously related pleurotomarians that we may specu-
late with considerable safety on the general nature of bellerophont
soft anatomy and perhaps even on its physiology and habits.

One may be quite confident that they were aspidobranchs with a
high degree of bilateral symmetry reflected in symmetrically paired
ctenidia, osphradia, hypobranchial glands, auricles, kidneys, and per-
haps even gonads. They probably crawled on the sea bottom on a
generalized gastropod foot. It seems probable that like other aspido-
branchs they fed chiefly on vegetable matter and were rhipidoglossate.
Nothing is known of the bellerophont operculum, if there was one.

Perhaps a diagrammatic restoration qf some of the more significant
soft parts with an interpretation of the course of the water currents
in the mantle cavity will save pages of words.

On figure 10 is shown a shell of a bellerophont species, Knightites
multicornutus Moore, 1941 (p. 153), with the soft parts restored in

Arrows show mantle cavity
currents.

Functional inhalant canols

Lae Sk
ae 1> Slit (exholont)
\ j 7
Left osphradium “Anus
~ A 2 ‘ 2 '
Left ctenidium e ight osphradium
~~ > ;
Tree “&, Right ctenidium
Sa
i aa ee — Slit- band
4
7
4
id
4
4
oa
Anterior protuberance of the parietal callus.
Pedal retractor musculature not shown.
|
Abandoned inhalant canals
eas ane
Osphradium ~~~ Left ctenidium
Ss 1 R Rees. ly Area of attachment of left
Slit (extdlont) ectum ~s pedal retractor muscle.

Functional. inhalant
NON Nya

canal
\
\

~~ Parietal Callus

Foot, with left pedal retractor muscle,
passing on the near side of the parietal

callus.
b

FIGURE IO

(See opposite page for explanation.)
54

NO. 13 PRIMITIVE FOSSIL GASTROPODS—KNIGHT 55

terms of the above interpretation. It is a modification of figure 7d in
Moore, 1941 (p. 158). I have abandoned the ideas expressed in
figures 7a-c as untenable in the light of more accurate knowledge of
the aerating currents in Haliotis than I then had. The extended
periodic, paired canals on each side of the slit and slitband in K. multt-
cornutus interested me very much. It occurred to me that they gave
a clue to the region on the mantle lip through which passed the cur-
rents of water that aerated and flushed out the mantle cavity. The
works of Yonge and Crofts on the aerating currents in various gas-
tropods including the pleurotomarian Haliotis seem to reinforce the
suggestion made by the canals of K. multicornutus, so that one can
infer the probable course of the principal water currents in that species
and probably in all generalized bellerophonts. This inferred circula-
tion is in all respects that of a prosobranch and seems a reasonable
approximation to the probable condition during life.

REFERENCES
Crorts, D. R.
1937. The development of Haliotis tuberculata, with special reference to
organogenesis during torsion. Philos. Trans. Roy. Soc. London,
ser. B, No. 552, vol. 228, pp. 219-268.
GarstTAnc, W.
1929. The origin and evolution of larval forms. British Assoc. Adv. Sci.,
Rep. 96th Meeting, pp. 77-98.
HowE tt, B. F., ET AL.
1944. Correlation of the Cambrian formations of North America. Bull.
Geol. Soc. Amer., vol. 55, pp. 993-1004.
KnicutT, J. Brookes.
1941. Paleozoic gastropod genotypes. Geol. Soc. Amer. Spec. Pap. 32.
1947a. Some new Cambrian bellerophont gastropods. Smithsonian Misc.
Coll., vol. 106, No. 17.

FIGURE I0

Two views of a diagrammatic restoration of Knightites multicornutus Moore
(Bellerophontacea, Bellerophontidae), approximately X 2. The shell is drawn
as though it were partly transparent so as to show some of the fleshy organs.
The arrows show the probable path of the principal aerating and cleansing
currents. Although most bellerophonts did not have inhalant canals as did
Knightites it is thought that the path of the currents and regions of their
entrance and exit were approximately the same as inferred for Knightites.
In living Haliotis where the details are known the paths are homologous in
every respect.

a, Dorsal view.

b, Left side view. The fibers of the left retractor muscle are shown in a

highly schematic fashion as though they anastomose into the muscles of the
foot (which are not shown).

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

1947b. Bellerophont muscle scars. Journ. Paleontol., vol. 21, pp. 264-267.
1948. Further new Cambrian bellerophont gastropods. Smithsonian Misc.
Coll., vol. 111, No. 3.
KosayAsHI, T.
1939. Restudy of Lorenz’s Raphistoma broggeri from Shantung, with a
note on Pelagiella. Jubilee publication in commemoration of Prof.
H. Yabe’s birthday.
LANKESTER, Ray.
1878. Mollusca. Encyl. Britt., 9th ed., vol. 16.
LInDsTROM, G.
1884. On the Silurian Gastropoda and Pteropoda of Gotland. Kong.
Svenska Vet.-Akad. Handl., vol. 19, No. 6.
LocHMAN, CHRISTINA.
1944. In Lochman and Duncan, Early Upper Cambrian faunas of central
Montana. Geol. Soc. Amer. Spec. Pap. 54.
Moors, R. C.
1941. Upper Pennsylvanian gastropods from Kansas. State Geol. Surv.
Kansas Bull. 38.
Naer, A.
1911. Studien zur generellen Morphologie der Mollusken. Teil 1. Uber
Torsion und Asymmetrie des Gastropoden. Ergebn. und Fortscher.
Zool., vol. 3, pp. 73-164.
PELSENEER, P.
1906. Mollusca. Jn Lankester, E. R., A Treatise on Zoology, pt. 5.
StmrotH, H.
1906. Jn Bronn, H. G., Klassen und Ordnungen des Tier-Reichs.

THIELE, J.
1931-1935. Handbuch der systemtischen Weichtierkunde.
WENz, W.
1938-1944. Gastropoda. Jn Schindewolf, O. H., Handb. Palaeozool., vol. 6,

Dt. 150-7.
1940. Ursprung und friithe Stammes-geschichte der Gastropoden. Arch.
Molluskunde, vol. 72, pp. I-10.
Woopwaprp, S. P.
1851-1856. A manual of the Mollusca, or rudimentary treatise of Recent
and fossil shells.
Yonce, C. M.
1939. On the mantle cavity and its contained organs in the Loricata
(Placophora). Quart. Journ. Micr. Sci., vol. 81, pp. 367-390.
1947. The pallial organs in the aspidobranch Gastropoda and their evolu-
tion throughout the Mollusca. Philos. Trans. Roy. Soc. London,
ser. B, vol. 232, pp. 443-518.

Just as this work reached page-proof stage, a copy of the “Traité de Paléon-
tologie,” published under the direction of Prof. Jean Piveteau, reached Wash-
ington (Traité de Paléontologie, vol. 2, 1952). The chapter on the Gastropoda
is by Dr. Geneviéve Termier (née Delpey) and Prof. Henri Termier of Algiers.
Since I have discussed Mme. Termier’s views on gastropods elsewhere (Geol.
Mag., vol. 83, pp. 280-284, 1946), I shall say nothing further here except to
reaffirm my almost complete disagreement.—J. B. K.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 27, NOS; ,PE

Coreospiridae

Cyrtolitidae
ES

PLATE I
1-3. Monoplacophora.
1. Scenella reticulata Billings. Lower Cambrian, Newfoundland. Left
side. Adult. Approx. * I. 2a, b. Helcitonella subruyosa (Orbigny).
Lower Cambrian, Troy, N. Y. a, Right side. b, View from above.
An example of the low, cup-shaped species. Approx. X I.
3a, b. Archaeophiala antiquissima (Hisinger). Upper Ordovician,
Sweden. a, Right side. b, View looking directly into the cup
of the shell showing the muscle scars (darkly pigmented) and
the “shadow scars.” This drawing was not made from the speci-
men but from photographs (Knight, 1941, pl. 3, fig. 3a-b). The
shell is shown a little too narrow posteriorly. Approx. X I.
4. Helcionella? sp. Middle Cambrian, Silver Peak Range, Nev.
An example of the high, narrow species of Helcionella?. Ap-
DUO
5-7. Bellerophontacea, Coreospiridae.
5. Oelandia acutacosta (Walcott). Lower Cambrian, Manuels Brook,
Conception Bay, Newfoundland. Left side view. Approx. X 4.
6. Oelandia sp. Middle Cambrian, Eagle, Alaska. Right side view.
Approx. X 5. 7a, b. Coreospira walcotti Knight. Lower Middle
Cambrian, Mount Stephen, British Columbia. a, View from above.
Note the total lack of an anal emargination. b, Right side view.
8,9. Bellerophontacea, Cyrtolitidae.
8a-d. Strepsodiscus major Knight. Upper Cambrian (Dresbachian),
Colorado. a, Anterior view showing V-shaped anal emargination.
b, Posterior view showing the slight asymmetry. c, Right side view.
d, Apertural view of specimen oriented on its side. Compare with
plate 2, fig. 7, Scaevogyra swesey!. Approx. X 1. ga-c. Cyrtolites
subplanus Ulrich. Middle Ordovician, Tennessee. a, Anterior view,
showing shallow V-shaped sinus. b, Apertural view from below.
c, Left side view. Approx. x I.
10, 11. Bellerophontacea, Sinuitidae.
toa, b. Sinuella minuta Knight. Upper Cambrian (Upper Dresbachian
or Lower Franconian), near Burnet, Tex. a, Anterior view. Note
U-shaped anal emargination. b, Left side view. Approx. X 10.
t1a, b. Simuites cancecllatus (Hall). Middle Ordovician (Trenton),
New York. a, Anterior view. Note U-shaped anal emargination.
b, Right side view. Approx. X I.
12, 13. Bellerophontacea, Bellerophontidae.
12a,b. Chalarostrepsis praecursor Knight. Upper Cambrian (Upper
Franconian or Trempealeauan). a, Anterior view. Note that the
anal emargination takes the form of a deep slit. b, Right side view.
Approx. * 1. 13a-c. Bellerophon sp. Mississippian, Indiana. a,
Right side view. b, Apertural view from below. c, Anterior view.
Note short slit. Approx. x I.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 217, NO 3 Ree

Pleurotomariacea

PLATE 2

1-6. Pleurotomariacea.

ta-b. Sinuopea sweeti (Whitfield). Upper Cambrian (Jordan member,
Trempealeauan), Wisconsin. a, Apertural view. b, Dorsal
view. Note the U-shaped anal emargination and compare it
with that of Sinuwites cancellatus (pl. 1, fig. 11a). Approx. I.
2. Schisopea typica (Ulrich and Bridge). Lower Ordovician
(Van Buren), Missouri. Note the deep V-shaped sinus.
Approx. X 1. 3. Loxoplocus (Lophospira) milleri (Miller).
Middle Ordovician (Trenton), New York. Note deep V-shaped
sinus generating a band. Approx. «14. 4. Entemnotrochus
adansoniana (Crosse and Fischer). Living, Caribbean Sea.
This is one of the species miscalled ‘‘Pleurotomaria” by neon-
tologists. Note the deep slit. Approx. & 1. 5. Phanerotrema
labrosum (Lindstrom) not Hall. Lower Silurian, Gotland. The
slit, which generates a band, is short. Approx. & 1. 6. Goniasma
sp. Pennsylvanian, Texas. Note the V-shaped sinus and the very
short slit. This is an example of the Murchisoniidae. Ap-
prox. < 14.

7-12. Macluritacea. All these are dextral and hyperstrophic. Hence the orien-
tation, actually the same as for other dextral forms, appears to be
up-side down. 7. Scaevogyra sweseyi Whitfield. Upper Cambrian
(St. Lawrence Member, Trempealeauan). Note the sharp, V-shaped
sinus culminating at the circumbilical carina. The sinus is the anal
emargination without a reasonable doubt. It should be noted here that
the thin shell with expanded aperture in Whitefield’s restoration is an
error. Wenz (1938, p. 239) grossly exaggerates the error. Approx. TI.
8. Lecanospira compacta (Salter). Lower Ordovician (Roubidouxian),
Quebec. Note the deep, V-shaped anal emargination and the circum-
bilical carina. Approx. 1. 9a-b. Onychochilus reticulatum Lind-
trom. Lower Silurian, Gotland. (After Lindstrom). a, A longitudinal
section. b, Apertural view. Note that the anal emargination is obscure
and that circumbilical ridge is rounded and troughlike. Approx. 2.
10. Matherella saratogensis (Miller). Upper Cambrian (Little Falls,
Trempealeauan), New York. As in Onychochilus the anal emargina-
tion is obscure and the circumbilical ridge rounded and troughlike.
Approx. <4. 11. Palliseria longwelli (Kirk). Middle Ordovician
(Chazyan), Nevada. The anal emargination is angular and culminates
at the angular circumbilical ridge. Approx. *& 4. (See also text figure
9, p. 41.) 12a-b. Maclurites logani Salter. Middle Ordovician (Black
River), Ontario. a, The operculum, inner surface. Note the attach-
ment rugosities of the left (1) and right (R) retractor muscles. As in
Nerita the attachment rugosity of the left muscle is an extended pro-
jection. b, The shell with the operculum in place, apertural view. The
umbilicus is narrower and the circumbilical ridge more rounded than
in some species. Note the counterclockwise spiral of the operculum, a
feature diagnostic of a dextral gastropod. Approx. X §.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 14

— Charles 3. and Mary Waux Walcott
Research Fund

|| | NEW AND UNUSUAL SPECIES OF
‘|| BRACHIOPODS FROM THE ARBUCKLE
| GROUP. IN OKLAHOMA

(Wits Four Piates)

BY
G. ARTHUR COOPER

Curator, Division of Invertebrate Paleontology and Paleobotany
Uv. ational Museum

(Pustication 4093)

i. “CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 23, 1952

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 14

Charles DB. and flary Waux Talcott
Research Fund

NEW AND UNUSUAL SPECIES OF
bRACGHiOPODs: FROM- THE. ARBUCKIEE
GROUP IN OKLAHOMA

(WitH Four PLates)

BY
G. ARTHUR COOPER

Curator, Division of Invertebrate Paleontology and Paleobotany
U. S. National Museum

(Pustication 4093)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 23, 1952

The Lord Baltimore Dress

BALTIMORE, MD., U. S. A.

Charles DB. and Mary Waux Walcott Research Fund

NEWrAND UNUSUAL SPECIES OF) BRACHIO-
FOODS FROM THE ARBUCKLE GROUP
IN OKLAHOMA
By G. ARTHUR COOPER

Curator, Division of Invertebrate Paleontology and Paleobotany
U. S. National Museum

(With Four PLates)

The brachiopods described herein resulted from collecting by Dr.
William E. Ham during the detailed mapping of the Arbuckle Moun-
tains. As might be expected, the great care taken by Dr. Ham during
the mapping to locate useful and easily recognizable fossils produced
a number of new species and others that had hitherto not been seen
in this region. Many of the occurrences produced beautifully silici-
fied specimens which make possible easy recovery of excellent study
material.

Since the appearance of Ulrich and Cooper’s study of the Ozarkian
and Canadian Brachiopoda, the stratigraphic sequence of the Ar-
buckle Mountains has been better defined by Decker (1939) and the
brachiopods can now be assigned to their proper formations. The
lists below include already known species and those described in this
paper.

Fort SILL FORMATION :

Billingsella corrugata Ulrich and Cooper.

Plectotrophia bridget Ulrich and Cooper.

laticosta Cooper, new species.

Mesonomia magna Cooper, new species.

SIGNAL MouNTAIN FORMATION :

Apheoorthis ornata Ulrich and Cooper.
oklahomensis Ulrich and Cooper.
platys Cooper, new species.

Billingsella rectangulata Cooper, new species.

Cymbithyris hami Cooper, new genus and species.

Fasciculina fasciculata Cooper, new genus and species.

Finkelnburgia auriculata Cooper, new species.

biconvexa Cooper, new species.
extensa Cooper, new species.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 14

Palaeostrophia cf. P. elax (Clark).
Glyptotrophia rotunda Cooper, new species.
McKeEnziz HILL FORMATION:

Finkelnburgia arbucklensis Cooper, new species.
cf. F. bellatula Ulrich and Cooper.
obesa Cloud.

Syntrophina campbelli (Walcott).

Tetralobula texana Ulrich and Cooper.

Coot CREEK FORMATION:
Finkelnburgia delicatula Cooper, new species.
subquadrata Cooper, new species.
Imbricatia lamellata Cooper, new genus and species.
Clarkella species.
Diaphelasma oklahomense Ulrich and Cooper.
Syntrophinella deckeri Cloud.

KINDBLADE FORMATION :

Finkelnburgia crassicostellata Cooper, new species.
cullisoni Ulrich and Cooper.
scemdioides Ulrich and Cooper.

Tritoechia delicatula Ulrich and Cooper.

subaequiradiata Ulrich and Cooper.
typica (Ulrich).

Oligorthis arbucklensis Ulrich and Cooper.

Diparelasma fasciculatwm Cooper, new species.

West SprING CREEK FORMATION:
Diparelasma costellatum Cooper, new species.
typicum Ulrich and Cooper.
Polytoechia subrotunda Ulrich and Cooper.
subcircularis Cooper, new species.
Pomatotrema magnum Ulrich and Cooper.
oklahomense Ulrich and Cooper.
transversum Ulrich and Cooper.
Syntrophopsis laevicula Ulrich and Cooper.
magna Ulrich and Cooper.
Finkelnburgia magna Ulrich and Cooper.
Tritoechia planodorsata Ulrich and Cooper.

Genus APHEOORTHIS Ulrich and Cooper, 1936
APHEOORTHIS PLATYS Cooper, new species

Plate 2B, figures 8-13

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

117

Shell small for the genus, wider than long, and subrectangular in
outline ; hinge forming the greatest width ; cardinal extremities slightly
auriculate. Sides nearly straight; anterior margin broadly rounded.

Surface fascicostellate.

Pedicle valve gently convex in lateral profile and with the greatest
curvature just anterior to the umbo; anterior profile broadly convex

NO. 14 BRACHIOPODS FROM OKLAHOMA—COOPER 3

but with the median region somewhat narrowly convex; beak small ;
umbo narrowly convex. Fold marked by a median and two lateral
fascicles, broad and low. Median region from umbo to anterior
margin moderately swollen ; flanks descending by a long, moderately
steep slope to the margins. Interior with strong dental plates but
only slightly thickened muscle area to form a spondylium discretum
or the incipient stages of a pseudospondylium.

Brachial valve slightly convex in lateral profile and broadly but
gently convex in anterior profile. Umbo smooth and slightly swollen ;
sulcus originating just anterior to umbo and widening and deepen-
ing to the anterior margin where it occupies one-third the valve width.
Flanks gently swollen; slopes to posterolateral extremities short and
gentle; cardinal extremities flattened. Interior with strong median
ridge, small simple cardinal process on a moderately thickened
notothyrial platform; brachiophores short and stout; adductor scars
strongly impressed.

Measurements in mm.

3 Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

TSN. NowanG737ay enc 9.0 e 9.8 10.0 2.2
Brachial valve,

(WESENaMinE NON Ti67 36D isaac ie ? 7.0 9.6 9.4 0.9

Types.—Holotype: U.S.N.M. No. 116737a; paratype: U.S.N.M.
No. 116736b.

Horizon and locality—Signal Mountain formation (50 feet below
the top) in Oklahoma, 2,100 feet north and 1,000 feet west of the
southeast corner of sec. 26, T. 1 S., R. 1 W., Murray County.

Discussion.—This species is characterized by a fairly flat brachial
valve, moderately convex pedicle valve and somewhat subdued orna-
mentation. This latter feature distinguishes this species from de-
scribed species. Its ornamentation is much more subdued than that
of A. ornata Ulrich and Cooper from the same formation.

Genus BILLINGSELLA Hall, 1892
BILLINGSELLA RECTANGULATA Cooper, new species
Plate 1A, figures I-15

Shell moderately large for the genus, wider than long, subrectan-
gular in outline. Hinge as wide as or slightly wider than the mid-
width; cardinal extremities slightly auriculate; sides nearly straight ;
anterior margin broadly rounded; surface somewhat fascicostellate,

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. P17

Pedicle valve nearly flat in the median region in lateral profile but
with the umbonal and anterior marginal regions somewhat narrowly
convex; anterior profile nearly flat to broadly and gently convex;
umbo narrowly convex, the convexity continued to about the valve
middle where it fades into the shell surface or slightly anterior to
the middle: anteromedian region flattened; anteromarginal region
gently but abruptly convex where a shallow, poorly defined sulcus
appears ; flanks bounding fold gently concave to nearly flat. Interarea
orthocline ; pseudodeltidium wide and strongly convex. Teeth strong,
buttressed by excess shell suggesting short dental plates; adductor
track moderately thickened anteriorly; pallial marks moderately
strong.

Brachial valve deeper than the pedicle one, gently convex in lateral
profile; broadly and moderately convex in anterior profile; umbo sul-
cate, sulcus wide and moderately deep to the valve middle but becom-
ing shallow and even, or almost so, with the flanks at the front margin.
Posterolateral areas flattened and with short steep slopes; flanks
bounding sulcus moderately swollen. Interior with short, stout
median ridge, lightly developed notothyrial platform and slender
cardinal process. Chilidium only moderately developed.

Measurements in mm.

Brachial Mid- Hinge Thick-

Length length width width ness

ULS MINNIS ING, MiOWAZE ooh cen odos 12.0 10.2 14.3 14.4 3.9
OKSINEIMIG ING MICA wo ais oer 13.5 II.0 14.8 14.9 4.3
URS UINIME INIGS IMG7AACd st oge ao oe 12.1 9.7 13.9 14.3 3.8

Types.—Holotype: U.S.N.M. No. 116722a; figured paratypes:
U.S.N.M. Nos. 116722b, d, e, h, i, k; unfigured paratypes: U.S.N.M.
Nos:/116722¢) 1, 2.4;

Horizon and locality—Signal Mountain formation (1-foot bed
of limestone 100 feet above the base) in Oklahoma, 1,700 feet east
and 1,300 feet north of the southwest corner of sec. 21, T. 1 S.,
R. 1 W., Murray County.

Discussion.—This species is characterized by its large size, rec-
tangular outline, and moderately short interarea. The specimens
illustrated are unusual in showing both valves in contact. Buillingsella
is generally loosely articulated and the valves fall apart after death.

Billingsella rectangulata is larger and squarer than B. coloradoensis
and is differently ornamented. It approaches B. corrugata Ulrich and
Cooper in size but lacks the concentric corrugations and acute cardinal
extremities of that species.

NO. 14 BRACHIOPODS FROM OKLAHOMA—COOPER 5

CYMBITHYRIS Cooper, new genus

Shell attaining a width of about a half inch, wider than long, and
with a wide hinge. Profile concavo-convex. Valves of unequal depth,
the pedicle valve having the greater depth. Surface multicostellate.

Pedicle valve with pseudodeltidium and foramen as in Buillingsella;
teeth small, dental plates absent ; musculature as in Bullingsella.

Brachial interior with small flat brachiophores like those of Billings-
ella; median ridge short and low; cardinal process a simple ridge.
Chilidium well developed.

Genotype—Cymbithyris hami Cooper, new species.

Discussion—The nearest relative to this genus is Billingsella. The
two genera have the same type of pseudodeltidium, teeth, and mus-
cular arrangement in the pedicle valve. They differ in that Cymbr-
thyris has a concave brachial valve. The brachiophores are like those
of Billingsella but other details of the brachial interior are obscure.
Cymbithyris is thus a lateral development from Bullingsella.

CYMBITHYRIS HAMI Cooper, new species
Plate 1B, figures 16-21

Shell subrectangular to quadrate in outline; hinge wider than mid-
width; cardinal extremities somewhat auriculate; sides oblique;
anterior margin narrowly rounded. Surface multicostellate.

Pedicle valve strongly convex in lateral profile; anterior profile
narrowly convex in the median region and with steep lateral slopes.
Umbo narrowly convex, the convexity extending to about the middle
where it is dissipated in the median swelling of the valve; median
region swollen, the broad swelling extending to the front margin.
Flanks moderately swollen and with long, moderately steep slopes to
the sides; posterolateral extremities somewhat flattened. Interarea
moderately long, orthocline.

Brachial valve fairly concave in anterior and lateral profiles ; umbo
sulcate, sulcus extending from umbo to front margin, expanding
anteriorly ; flanks bounding sulcus moderately concave.

Measurements in mm.

c Brachial Mid- Hinge ; Thick-
Pedicle valves, Length length width width Height ness

U.S.N.M. No. 116724a.... 12.4 ? 13.7 16.6 4.5 ?

U.S.N.M. No. 116724b.... 12.7 f 14.3 16.5 3.8 ie

U.S.N.M. No. 116724d.... 9.5 8.2 ? 2 ? 2.2

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. lly

Types.—Holotype: U.S.N.M. No. 116724d; figured paratypes:
U.S.N.M. Nos. 116724 a-c, e-f.

Horizon and locality.—Signal Mountain formation (limestone one
foot thick 126 feet above the base) in Oklahoma, 1,700 feet west and
600 feet south of the northeast corner of sec. 28, T. 1 S., R. 1 W.,
Murray County.

Discussion—No other species of this genus is known to which
C. hami can be compared.

Genus MESONOMIA Ulrich and Cooper, 1936
MESONOMIA MAGNA Cooper, new species
Plate 1D, figures 27-34

Shell large for the genus, subrectangular in outline with the hinge
forming the widest part; cardinal extremities auriculate ; sides nearly
straight ; anterior margin broadly rounded. Anterior commissure
uniplicate. Ornamentation consisting of costellae of varying sizes
alternating with each other.

Pedicle valve unevenly convex in lateral profile and with the
maximum convexity in the umbonal region; anterior profile broadly
convex. Umbo narrowly swollen and forming a narrow fold to about
the valve middle; anterior part of the median portion bent fairly
abruptly in the direction of the brachial valve and forming a short,
narrowly convex tongue; flanks bounding sulcus gently swollen;
slopes to the posterolateral extremities short and gentle. Interior
with small teeth and moderately thickened adductor track.

Brachial valve gently convex in lateral profile and broadly but
gently convex in anterior profile; umbo sulcate, sulcus shallow and
continued anteriorly to about the middle where it suddenly reverses
to form a moderately broad fold; flanks gently swollen; interior with
short, delicate brachiophores supported by shallow plates that unite
under the thin cardinal process. Median ridge short and delicate.

Measurements in mm.

< Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

U.S.N.M. No. 116604a..... 12.3 ? 15.0 15.8 Be
Brachial valves,

U.S.N.M. No. 116694b..... ? 10.7 14.8 17.8 2.9

U.S.N.M. No. 116604c..... ? II.I 15.8 172 ZY,

Types——Holotype: U.S.N.M. No. 116694a; figured paratypes:
U.S.N.M. Nos. 116694b-d.

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 7.

Horizon and locality—Fort Sill formation (179 feet above the
base) in Oklahoma, 1,000 feet south of the north quarter corner of
Seen 20. Dat o. Rt Ey Murray ‘County.

Discussion.—This species is distinguished by its large size, gently
sulcate pedicle valve, and the nature of the ornamentation. The
species is most like M. iophon (Walcott) from the Mons formation
of Alberta but is much larger and with less pronounced fold and
sulcus. The ornamentation of M. iophon is stronger than that of the
Arbuckle species and is not so strongly differentiated into strong and
fine costellae.

FASCICULINA Cooper, new genus

Exterior somewhat resembling Apheoorthis and with unequally
convex valves, the pedicle valve having the greater depth. Anterior
commissure broadly sulcate; surface fascicostellate.

Pedicle valve with swollen median region serving as a fold; teeth
small; dental plates well developed; muscle field orthoid and with
the adductor track more or less elevated to form a pseudospondylium.

Brachial valve with sulcus originating at the beak and usually
bounded by two strong costellae; cardinalia delicate; brachiophores
short, small; sockets formed by small fulcral plates; brachiophore
supports nearly erect to moderately oblique, attached directly to the
floor of the valve but more or less joined by a callosity between them ;
cardinal process a simple ridge when present. Median ridge formed
by inner swelling corresponding to sulcus; adductor callosities thick
in old shells.

Genotype.—Orthis desmopleura Meek (as redefined by Ulrich and
Cooper, Geol. Soc. Amer. Spec. Pap. 13, p. 131, 1938).

Discussion—This genus can be recognized by its fasciculate ex-
terior, usually weakly developed pseudospondylium, and discrete
brachiophore plates. It may be easily confused with Finkelnburgia
when the brachiophore plates are strongly oblique, but it differs from
that genus in having a much stronger sulcus, one that extends from
beak to anterior margin, much stronger ornamentation and generally
weaker development of the pseudospondylium.

Fasciculina is like Orusia internally but is much differently orna-
mented. Furthermore in that genus the dental plates are short and
no trace of a pseudospondylium has been seen.

Other species besides the genotype assigned to Fasciculina are
Eoorthis wichitaensis Walcott and possibly E. indianola Walcott.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, I1I7

FASCICULINA FASCICULATA Cooper, new species
Plate 2A, figures 1-7

Small, subquadrate in outline, wider than long, and with the hinge
forming the widest part; sides slightly oblique; anterior margin
broadly rounded; cardinal extremities slightly auriculate. Surface
fascicostellate.

Pedicle valve evenly and moderately convex in lateral profile and
with the maximum convexity at about the middle; anterior profile
fairly strongly and broadly convex; umbo narrowly convex, the con-
vexity extended to the anterior margin as a low and broad fold
having a median costella at the umbo and extending to the margin
but with a broad fascicle implanted on each side of it. Flanks gently
swollen and separated from the fold by a strong costella on each side
giving the appearance of a fold within a broad median sulcus. Pseudo-
spondylium moderately thickened and with the adductor track slightly
elevated.

Brachial valve gently convex in lateral profile and broadly and
gently convex in anterior profile ; sulcus originating at the umbo and
widening rapidly to the front margin where it occupies about half the
width; posterior of sulcus marked by two costellae which bound a
deep median sulcus within the broad sulcus; broad sulcus bounded by
a strong costella on each side, and these oppose the depressions bound-
ing the pedicle fold and the deeper inner sulcus opposes the fold within
the pedicle depression. Flanks bounding sulcus, narrow, gently
swollen. Interior with short delicate brachiophores and recumbent
brachiophore plates.

Measurements in mm.

; Brachial Mid- Hinge Thick-
Pedicle valves, Length length width width ness

W.SiN MING? aie7s8an.e oe 6.4 ? 7.9 7.9 plus 2.4

WES DINIIMI, INKen iiiloyst8}0), on goons Fo ? 8.6 8.9 2.3
Brachial valve,

WES SNe MesNomrro7a8dienancs ? 5.7 7.8 8.1 7

Types——Holotype: U.S.N.M. No. 116738e; figured paratypes:
U.S.N.M. Nos. 116738b-d; unfigured paratype: U.S.N.M. No.
116738a.

Horizon and locality.—Signal Mountain formation (go feet above
the base) in Oklahoma, in the SESSW4NE4 sec. 16, T. 1 S., R. 1 W.,
1,200 feet west of the east quarter corner, Murray County.

Discussion.—This species can be recognized by its strong costellae,
alate cardinal extremities, fairly deep sulcus, and fairly well-developed

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 9

pseudospondylium. It most resembles F. desmopleura (Meek) in its
ornamentation but differs in its interior, the brachiophore supporting
plates of the Arbuckle shell being more oblique and joined by a callus
on the floor of the valve. Furthermore F. fasciculata has a moderately
developed pseudospondylium whereas that of F. desmopleura is not
strongly thickened.

This species differs from F’. wichitaensis in being much smaller,
having a more pronounced pedicle fold and stronger costellae.

Genus FINKELNBURGIA Walcott, 1905
FINKELNBURGIA ARBUCKLENSIS Cooper, new species
Plate 3H, figures 33-39

Shell of about medium size for the genus, wider than long; sub-
rectangular in outline; sides nearly straight ; hinge equal to or slightly
narrower than the midwidth; cardinal extremities slightly obtuse or
forming a right angle. Multicostellate with a few strong costellae
standing out in a mat of fine ones.

Pedicle valve moderately convex in lateral profile but with the
maximum curvature in the umbonal region; broadly and strongly
convex in anterior profile; beak prominent ; umbo and median regions
swollen ; flanks full with short steep slopes to the margins. Delthyrial
cavity wide and deep; pseudospondylium with well-developed dental
plates, narrow or broad adductor track but short median ridge. Pallial
trunks wide and deeply impressed.

Brachial valve shallower than the pedicle valve, with gentle con-
vexity in lateral profile and broadly convex in anterior profile. Sulcus
poorly defined, shallow, extending from umbo to anterior margin;
flanks gently swollen but median region somewhat flattened. Lateral
slopes short and gentle. Interior with broad and short notothyrial
cavity; adductor callosities not strongly developed but six pallial
trunks deeply impressed.

Measurements in mm.

: Brachial Mid- Hinge Thick-
Pedicle valves, Length length width width ness

ULES EINEIMIS INS TG 7/E oegoooar 8.9 ? 11.3 9.6 2.5

SMe ING: MIGZAUD ss... 8.0 ? 10.3 10.3 22
Brachial valves,

OESENEMeINiowimnG7AlCa.s 14) 1. ? 8.8 12.0 10.5 2.3

W2SsNeMeeNow 6740s... =... ? 9.2 12.1 11.8 2.3

Types —U.S.N.M. No. 116741b; figured paratypes: U.S.N.M.
Nos. 116741a, c, d.

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

Horizon and locality—McKenzie Hill formation (237 feet above
the base) in Oklahoma, 1,500 feet east and 2,000 feet south of the
northwest corner of sec. 28, T. 1 S., R. 1 W., Murray County.

Discussion.—This species is characterized by subrectangular out-
lines nearly rectangular or slightly obtuse cardinal extremities, poorly
defined sulcus and closely crowded costellae. The species suggests
F, bellatula Ulrich and Cooper but that is a smaller and somewhat
less transverse shell with stronger costellae and more delicate pseudo-
spondylium. Finkelnburgia arbucklensis is simliar to F. hellert Cloud
but differs in being less transverse and in having a finer ornament,
stouter cardinalia, and a narrower pseudospondylium.

FINKELNBURGIA AURICULATA Cooper, new species
Plate 2F, figures 29-38

Shell small, wider than long, and with the hinge wider than the
midwidth ; cardinal extremities mucronate; sides gently oblique; an-
terior margin broadly rounded; surface fascicostellate.

Pedicle valve deeper than the brachial valve; strongly convex in
lateral profile; anterior profile strongly arched; umbo and median
region strongly swollen; flanks swollen and with steep slopes to the
margins. Interarea short, curved, apsacline. Interior with a deep
pseudospondylium but with the adductor track only moderately
thickened.

Brachial valve strongly convex in lateral profile; anterior profile
strong and broadly convex and with steep lateral slopes. Umbo sul-
cate; sulcus widening and deepening anteriorly to the front margin.
Flanks bounding sulcus fairly strongly swollen; brachiophores and
supporting plates stout.

Measurements in min.

= Brachial Mid- Hinge Thick-
Length; length width width ness
CESN. MM: Not mioyaita necscasns ice er 6.1 cy 7.4 7.9 4.7
Pedicle valve,
WESINEM eI NositG7aiteneeene eee 37) ? 7.0 8.0 20

Types—Holotype: U.S.N.M. No. 116731a; figured paratypes:
U.S.N.M. Nos. 116731c-f£; unfigured paratype: U.S.N.M. No.
116731b.

Horizon and locality.—Signal Mountain formation (6-inch bed of
limestone 150 feet above the base) in Oklahoma, 300 feet west and
1,150 feet south of the northeast corner of sec. 28, T. 1 S., R. 1 W.,
Murray County.

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER Wa

Discussion.—This species is characterized by rectangular outline,
strongly and subequally convex valves, and the sharp auriculation of
the cardinal extremities. In the latter respect the species resembles
F, finkelnburgt but it is much smaller and has straighter lateral margins
and less extended cardinal extremities.

FINKELNBURGIA cf. F. BELLATULA Ulrich and Cooper
Plate 3G, figures 28-32

Finkelnburgia bellatula UtricH AND Cooper, Journ. Paleontol., vol. 10, No. 7,
p. 622, 1936; Geol. Soc. Amer. Spec. Pap. 13, p. 134, pl. 25C, figs. 9, 10,
13-28, 1938.

Shell of about medium size for the genus, roundly subelliptical in
outline with the hinge shorter than the midwidth which is the widest
part; sides well rounded; anterior margin broadly rounded; cardinal
extremities obtuse. Multicostellate, costellae of different sizes.

Pedicle valve gently convex in lateral profile but moderately convex
in anterior profile; beak moderately protruberant; umbo narrowly
swollen; entire valve fairly strongly inflated and with long convex
lateral slopes. Interior with pseudospondylium short and weakly
developed.

Brachial valve less deep than the pedicle valve, moderately convex
in lateral profile; broadly and moderately convex in anterior profile ;
median regions inflated but posterior and median portion marked
by a shallow and narrow sulcus which does not reach the margin.
Anteromedian region swollen and conforming with the convexity of
the rest of the valve; lateral slopes short and steep; interior with
thick, erect brachiophore supports and thick cardinal process. Ad-
ductor platforms moderately developed.

Measurements in mm.

: Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

WES NUM GINO: WETG747 a. cele ccs 5 8.4 ? 10.1 7.0 2.2
Brachial valve,

RES Ma ING: E6747)... vss ? 7.6 9.8 6.8 2.1

Hypotypes.—U.S.N.M. Nos. 116747, b.

Horizon and locality—McKenzie Hill formation (10 feet below
the top) in Oklahoma, 2,200 feet north and 1,250 feet west of the
southeast corner of sec. 7, T. 1 S., R. 1 E., Murray County.

Discussion—The specimens here assigned to F. bellatula agree in
outline and profiles and the ornamentation is well differentiated as

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

in the type specimens. Furthermore the interiors agree in having a
delicate pseudospondylium and the adductor platforms confined to
the rear part of the shell.

FINKELNBURGIA BICONVEXA Cooper, new species
Plate 2E, figures 22-28

Shell of about medium size for the genus, wider than long, sub-
rectangular in outline; hinge about equal to midwidth; cardinal ex-
tremities slightly auriculate; sides nearly straight; anterior margin
broadly rounded; surface multicostellate, costellae crowded and of
different sizes.

Pedicle valve deeper than the brachial valve, strongly convex in
lateral profile; strongly convex in anterior profile and with long,
steep, lateral slopes. Umbo, median region, and flanks strongly inflated.
Pseudospondylium broad and low; adductor track thickened ; median
ridge short and thick.

Brachial valve moderately convex in lateral profile but broadly and
strongly convex in anterior profile; beak prominent; umbo somewhat
broadly and strongly swollen, the swelling continued anteriorly to
the margin; anterior third somewhat flattened to approximate a
median sulcus or definitely sulcate, the sulcus broad and shallow;
flanks somewhat swollen and with short steep slopes to the margins.
Interior with large and thick cardinalia; adductor platforms not
developed.

Measurements in mm.

$ Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

WS. NMA NOs 31673382 seat 8.1? ? 0.3 8.6 2.9
Brachial valve,

WESIN. Me INO. -216733t asec ? 6.8 0.3 8.6 2.3

Types.—Holotype: U.S.N.M. No. 116733a; figured paratypes:
U.S.N.M. Nos. 116733¢, e, £; unfigured paratypes: U.S.N.M. Nos.
116733b, d.

Horizon and locality —Signal Mountain formation (218 feet above
the base) in Oklahoma, 500 feet west and 750 feet south of the north-
east corner of sec. 28, T. 1 S., R. 1 W., Murray County.

Discussion.—This species is characterized by strongly convex
valves, closely crowded costellae, and a thick pseudospondylium. It
is quite unlike other known Upper Cambrian Finkelnburgias except
F, osceola which is larger and differently shaped, and F. newtonensis

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 13

(Weller) which is much smaller, less convex, and differently orna-
mented.

Of Canadian species this one is most like F. wemplei but differs
in greater convexity, particularly of the brachial valve, and in having
slight auriculations on the brachial valve.

FINKELNBURGIA CRASSICOSTELLATA Cooper, new species
Plate 1C, figures 22-26

Shells of intermediate size for the genus, wider than long, sub-
rectangular in outline and with the hinge narrower than the maximum
width which is at the middle; cardinal extremities obtuse; sides
rounded; anterior margin broadly rounded; surface strongly cos-
tellate, costellae of unequal size.

Pedicle valve gently convex in lateral profile and with the most
curvature just anterior to the middle; anterior profile moderately and
broadly convex, umbo broadly swollen; median region gently convex ;
anterior half somewhat swollen and curved toward the brachial valve ;
flanks moderately swollen and with short, steep lateral slopes. Pseudo-
spondylium broad and poorly developed ; median ridge low and thick.

Brachial valve much less deep than the pedicle valve, gently convex
in lateral and anterior profiles; umbo and median region gently
swollen; sides gently swollen and with long, gentle, lateral slopes;
interior with broadly recumbent brachiophore plates, and strong ad-
ductor callosities.

Measurements in mm.

z Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

U.S.NEM: No) 5n6726a./4n0.... 8.5 ? 11.0 8.8 2.5
Brachial valve,

UES.N.M.) Nos. 1167266. 90. : 7.5 10.9 10.2 1.8

Types——Holotype: U.S.N.M. No. 116726a; figured paratypes:
U.S.N.M. Nos. 116726b, c.

Horizon and locality.—Kindblade formation in Oklahoma, in the
center NES W3 sec. 29, T. 1 S., R. 1 W., Murray County.

Discussion—This species is characterized by strong costellae, a
shallow brachial valve having large adductor platforms extending to
about the middle and a fairly deep pedicle valve with broad, weakly
developed pseudospondylium. The species suggests a small F. magna
but it attains only about half the size of that species and also differs
in the strength of its costellae. The interior of both valves is not so

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

extravagantly developed as that of F. magna, but the structures are
of the same broad character.

FINKELNBURGIA DELICATULA Cooper, new species
Plate 2D, figures 16-21

Small for the genus, wider than long with the hinge less than or
about equal to the midwidth; cardinal extremities slightly obtuse or
approximating a right angle; sides nearly straight; anterior margin
broadly rounded; surface finely and closely costellate, but with scat-
tered strong costellae.

Pedicle valve moderately convex in lateral profile and with the
maximum convexity in the posterior third; anterior profile strongly
convex ; umbonal and median regions swollen ; flanks descending mod-
erately steeply to the margins; anterior somewhat flattened ; interior
with well-defined pseudospondylium with thickened and elongated
adductor track supported by a short stout median ridge.

Brachial valve not so deep as the pedicle valve and moderately con-
vex in lateral profile, broadly and moderately convex in anterior
profile. Umbo sulcate; sulcus narrow and shallow, defined from umbo
to anterior margin; flanks swollen and with short, moderately steep
lateral slopes. Interior with moderately well-developed adductor
platforms.

Measurements in mm.

. Brachial Mid- Hinge Thick-
Pedicle valve, Length length width wicth ness

WE SsNEME NOs 1107340 erase 7.6 ? 9.0 8.8 2.5
Brachial valves,

WWESSINGIMI, INGE MiGyuniag ao sunen ? 6.1 8.3 78 2.0

WES. Neve NON mT6734he ese se ? 5.9 8.4 7.3 17

Types.—Holotype: U.S.N.M. No. 116734a; figured paratypes:
U.S.N.M. Nos. 116734f, g; unfigured paratypes: U.S.N.M. Nos.
116734b-e, h.

Horizon and locality—Upper Cool Creek formation in Oklahoma,
one mile southeast of the windmill, center of sec. 5, T.2S., R. 1 E.,
Murray County.

Discussion.—Small size, fine costellae, subrectangular outline, and
subauriculate cardinal extremities characterize this species. It is
suggestive of F. bellatula in outline but has a somewhat wider hinge
with definite auriculations and somewhat finer ornamentation. The
interiors of the two species show differences. The muscle platforms
of the brachial valve of F. delicatula occupy more of the interior than

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 15

those of F. bellatula and the pseudospondylium of the pedicle valve
is stouter than that of F. bellatula.

FINKELNBURGIA EXTENSA Cooper, new species
Plate 2G, figures 39-51

Shell of about medium size for the genus, width of hinge equal to
about twice the shell length; cardinal extremities mucronate ; lateral
margins concave just anterior to the cardinal extremity; anterior
margin broadly rounded. Surface marked by crowded costellae of
unequal size, the larger ones in the minority, but numbering three or
four to a millimeter at the front margin.

Pedicle valve unevenly convex in lateral profile, with the maximum
convexity just anterior to the umbo; anterior profile narrowly convex
in the median region and with abrupt slopes steep on the flanks of
the median area but gentle to the margins. Beak prominent and ele-
vated; umbo narrow and convex; median region strongly swollen
from umbo to anterior margin; interior with pseudospondylium hav-
ing an extravagantly thickened adductor and median ridge; pallial
trunks deeply entrenched.

Brachial valve slightly less deep than the pedicle valve and having
a moderately convex lateral profile in which the maximum convexity
is located just anterior to the umbo; anterior profile broadly and
moderately convex. Sulcus narrow and shallow, originating on the
umbo and extending to the anterior margin, widening only slightly in
its passage anteriorly ; flanks bounding sulcus fairly strongly swollen ;
lateral slopes short, moderately steep. Interior with deep sockets,
short and blunt brachiophores and strong, thick brachiophore plates.
Cardinal process thin and delicate. Adductor platforms not strongly
thickened.

Measurements in mm.

3 Brachial Mid- Hinge Thick-
Pedicle valves, Length length width width ness
LES INEMG, Noe TIG728D.... ..2'55- 6.8 ? 8.8 13.3 2.9
WLSaINGIMIG ING, Misra ooas 8.9 ? 12.2 19+ 4.0
WESINEMNone16728hin. aoeeis- 7.2 ? 7.0 12.0 2.6
Brachial valves,
UESLINEIMIS IN\e, Tileyzsh aes ae abe ? 6.3 8.1 II.0 2.1
WSN ING. 21672844 nits... P 6.5 9.1 12.7 22
(WASSNEM Now t10728ie ae. . ee ? 5.9 8.7 13.4 1.9

Types.—Holotype: U.S.N.M. No. 116728b; figured paratypes:
U.S.N.M. No. 116728a, d-g, j; unfigured paratypes: U.S.N.M. Nos.
116728c, h, i, k.

Horizon and locality—Signal Mountain (90 feet above the base)

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

in Oklahoma, SES4SW4NE4 sec. 16, T. 1 S., R. 1 W., 1,200 feet west
of the east quarter corner, Murray County; 70 feet above the base,
2,300 feet west and 1,700 feet north of the southeast corner of sec. 8,
T.1S., R. 1 W., Murray County.

Discussion—This species is unusual for the great extension of its
cardinal extremities and the deep impression of the pallial channels
on the inside of the valves. It is most like the type species of Finkeln-
burgia, F. finkelnburgi Walcott, in the great extension of the cardinal
extremities but is more extreme in this respect than any figured
specimens of that species. Finkelnburgia extensa differs from F.
finkelnburgi also in having finer ornamentation.

FINKELNBURGIA OBESA Cloud
Plate 3F, figures 20-27

Finkelnburgia obesa CLoup, Bull. Mus. Comp. Zool., vol. 100, No. 5, p. 458,
pl. 1, figs. 8-18, 10948.

Shell of intermediate size for the genus, wider than long, suboval
in outline with the hinge narrower than the greatest width which is
just anterior to the middle; cardinal extremities obtuse; sides
rounded; anterior margin broadly rounded. Surface multicostellate.

Pedicle valve moderately convex in lateral profile but with the
maximum convexity in the umbonal region. Anterior profile broadly
convex; beak prominent ; umbo somewhat narrowly swollen but the
swelling not continued to the middle; median region and flanks gently
swollen ; anterior half somewhat depressed to form a poorly defined |
sulcus. Interior with prominent and well-developed pseudospondylium
having an elevated adductor track and a moderately high median
septum.

Brachial valve gently convex in lateral profile and not having as
much depth as the pedicle valve; anterior profile broadly and gently
convex. Umbonal, median and flank regions swollen; lateral slopes
short and steep. Interior with brachiophore plates stout and well-
defined ; cardinal process a thin ridge ; adductor platforms moderately
developed.

Measurements in mm.

: Brachial Mid- © Hinge —Thick-
Pedicle valves, Length length width width ness

WESINE MED Noyetro7Asaesneseer 9.5 & 11.4 6.9 2.8

OOSEINEINIS ING) WHGVZIE Do oo oosoe 9.3 ? 10.6 8.6 27
Brachial valves,

WESINEIMI ING) TMGVAIC55 Soo0ec & 10.4 11.6 8.7 3.0

WWESBINGIMG INK}, Viale HslS o Gooes fy 8.9 TI.5 8.0 2.4

WESAN:M: MNo. sm6745e.. .5-25- ? 8.4 10.5 72 23

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 17

Types.—Figured hypotypes: U.S.N.M. Nos. 116745b-e; unfigured
hypotype: U.S.N.M. No. 116745a.

Horizon and locality—McKenzie Hill formation (35 feet below
the top) in Oklahoma, 2,200 feet east and 2,500 feet north of the
southwest corner of sec. 2, T, 2 S., R. 1 E., Murray County.

Discussion.—Comparison of the McKenzie Hill specimens with
Cloud’s types of F. obesa indicates the exterior form to be very close.
Comparison of the ornamentation is not possible in detail because the
specimens from Oklahoma and Texas are badly worn. Those of the
Arbuckle species were undoubtedly considerably water-worn before
silicification. Details of the interior of the Texas species are not well
preserved, but those that can be seen show close resemblance except
for the fact that the Oklahoma specimens are somewhat thicker.

FINKELNBURGIA SUBQUADRATA Cooper, new species
Plate 31, figures 40-45

Shell large for the genus; wider than long and with the hinge
slightly wider than the midwidth ; cardinal extremities slightly auricu-
late ; sides gently rounded; anterior margin broadly rounded. Surface
strongly costellate, costellae of unequal size.

Pedicle valve considerably deeper than the brachial valve, strongly
convex in lateral profile and with the maximum convexity in the
umbonal region and just anterior thereto; anterior profile moderately
and broadly convex. Beak small, slightly protruberant. Umbo
swollen ; median region from umbo to anterior margin swollen ; flanks
swollen and with short steep slopes to the margins. Pseudospondylium
broad and with a long tonguelike adductor track ; median ridge short
and thick. Pallial sinuses (vascula media) wide and strongly
divergent.

Brachial valve moderately convex in lateral profile; broadly and
moderately convex in anterior profile. Umbo marked by a short,
shallow sulcus; umbo and median region swollen, the swelling re-
flected in the flanks; lateral slopes moderately short and moderately
gentle. Interior with small notothyrial chamber bounded by short
brachiophore plates. Adductor callosities modestly developed.

Measurements in mm.

, Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness
U.S.N.M. No. 116743a.... 12.8 e 15.5 14.7 plus 4.0

Brachial valve,
WES NEM No: 116743b. 0. ~ 10.2 13.7 13.6 3.0

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Types.—Holotype: U.S.N.M. No. 116743a; paratype: U.S.N.M.
No. 116743b.

Horizon and locality.—Cool Creek formation in Oklahoma, 900
feet west and 1,450 feet north of the southeast corner of sec. 3, T.
2S.,R.1 W., Murray County.

Discussion.—This species can be recognized by its large size, sub-
quadrate outline, strong costellae, thick and elevated pseudospon-
dylium, and the broad, divergent pallial marks in the pedicle valve.
Finkelnburgia subquadrata most strongly resembles F. virginica but
is somewhat larger, is more quadrate, and has more distant costellae.
Inside the brachial valve the adductor platforms of F. virginica are
scarcely developed but in the Arbuckle shell they are strong. Inside
the brachial valve the deep and wide vascula media are a feature
peculiar to the Oklahoma species.

Genus PLECTOTROPHIA Ulrich and Cooper, 1936
PLECTOTROPHIA LATICOSTA Cooper, new species
Plate 4F, figures 23-35

Shell of about usual size for the genus, wider than long, subrec-
tangular in outline; sides gently oblique to gently rounded; hinge
slightly wider or slightly less than the midwidth depending on age;
cardinal extremities slightly obtuse or slightly auriculate; anterior
commissure gently uniplicate; surface marked by distant, rounded,
elevated costellae crossed by prominent concentric lines of growth.

Pedicle valve unevenly convex in lateral profile and with the maxi-
mum convexity between the umbo and the middle, the anterior half
flattened. Anterior profile with the median region somewhat nar-
rowly convex and with steep sides. Beak small, umbo narrowly
convex; posteromedian region swollen and with steep slopes to the
cardinal extremities. Sulcus originating just anterior to the umbo,
deepening and widening near the middle. Anterior tongue short and
bluntly rounded. Interior with long and narrow pseudospondylium
tapering anteriorly to a sharp point ; median septum only slightly de-
veloped. Pallial marks not deeply impressed.

Brachial valve gently convex in lateral profile and with the maxi-
mum depth located just anterior to the umbo; anterior profile broadly,
fairly evenly and moderately convex. Beak small, umbo somewhat
narrowly swollen; median region broadly and strongly swollen; pos-
terior marked by a short, shallow depression just anterior to the
umbo, which is followed by the fold; fold originating just posterior
to the middle, widening anteriorly, low and broadly rounded through-

NO. 14 BRACHIOPODS FROM OKLAHOMA—COOPER 19
out. Flanks bounding fold gently swollen and with long gentle slopes
to the margins. Interior with short brachiophores supported by short

convergent supporting plates.

Measurements in mm.

Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

UPS NEMeuNoOs T7795 Tde.., 462116 8.5 ? 10.9 med 2.2?
Brachial valve,

Wes NeM= Nosmr7O5ibe essen. ? 8.1 12.7 II.4 a7

WES ENG Mi INion mI7OSITE. face's «le ? 7.6 II.5 10.2? 2.3

Types—Holotype: U.S.N.M. No. 117951g; figured paratypes:
U.S.N.M. Nos. 117951a-f; unfigured paratypes: U.S.N.M. Nos.
11795th-k.

Horizon and locality —Fort Sill formation (150 feet above the base)
in Oklahoma, 2,500 feet east and 2,200 feet north of the southwest
corner of sec. 26, T. 2 S., R. 4 E., South Ranch, Johnston County.

Discussion.—This species is especially characterized by its strong
costellae, the low and broad fold, and the short blunt tongue of the
pedicle valve. When compared with P. bridgei the Oklahoma speci-
mens have a much less-elevated and less-carinate fold, a shallower
sulcus on the pedicle valve, and a much shorter, blunter tongue on
the pedicle valve. Inside the pedicle valve the pseudospondylium of
P. bridgei is more elevated, has subparallel sides and a narrowly
rounded front, whereas the same structure in P. laticosta is long,
slender, tapers to a point, and is low throughout its extent.

Genus TETRALOBULA Ulrich and Cooper, 1936
TETRALOBULA TEXANA Ulrich and Cooper
Plate 4A, figures 1-7

Tetralobula texana ULricH and Cooper, Geol. Soc. Amer. Spec. Pap. 13, p. 200,
pl. 43B, figs. 17-22, 1938.—Ctoup, Bull. Mus. Comp. Zool., vol. 100, No. 5,
p. 462, pl. 2, figs. 15-16, 21, 22, 1948.

A few well-preserved specimens of this species are illustrated to
show the occurrence of this easily recognized genus in Oklahoma.
The species is characterized by its low fold which is located at the
anterior margin only. In the pedicle valve the sulcus is broad and
shallow and the tongue broad and rounded. In young specimens little
trace of the fold can be detected.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. Lily

Measurements in mm.

E Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

UESUN MeINoe 1r7O50en eee 8.7 ? II.0 8.5 3.0
Brachial valve,

USN. Me Norarzosone ote s ? 7.4 9.8 ES 2.2

Figured specimens.—U.S.N.M. Nos. 117950a-f.

Horizon and locality—McKenzie Hill formation (about 250 feet
above the base) in Oklahoma, 2,100 feet west and 1,500 feet north of
the southeast corner of sec. 36, T. 1 S., R. 1 E., Murray County.

Genus PALAEOSTROPHIA Ulrich and Cooper, 1936
PALAEOSTROPHIA cf. P. ELAX (Clark)
Plate 3A, figures I, 2

Syntrophia elax CiarK, Bull. Amer. Paleontol., vol. 10, No. 41, p. 15, pl. 3,
fig. I, 1924.

Palaeostrophia elax (Clark) UtricuH and Cooper, Geol. Soc. Amer. Spec.
Pap. 13,.p. 105, pl. 4rA, figs: 1-11, 15, 1038:

Shell of about usual size for the genus, represented by three frag-
mentary specimens. Pedicle valve with wide and shallow sulcus
occupying most of anterior region. Brachial valve with a broadly
rounded fold. Interior with characteristic erect brachiophore support-
ing plates and elongate adductor tracks.

Figured specimens.—U.S.N.M. Nos. 116751a, b.

Horizon and locality—Signal Mountain formation (63 feet below
the top) in Oklahoma, 1,500 feet west and 1,900 feet north of the
southeast corner of sec. 6, T. 2 S., R. 2 E., Murray County.

Genus GLYPTOTROPHIA Ulrich and Cooper, 1936
GLYPTOTROPHIA ROTUNDA Cooper, new species
Plate 2C, figures 14, 15

Small, transversely but broadly elliptical in outline; sides rounded ;
anterior commissure strongly uniplicate ; surface marked by fine cos-
tellae cancellated by strong concentric lamellae.

Pedicle valve gently convex in lateral profile and strongly but
broadly convex in anterior profile. Beak projecting ; umbo narrowly
rounded; median region swollen. Sulcus originating just posterior
to the middle, deepening anteriorly but shallow throughout its length.
Tongue short and narrowly rounded. Flanks swollen near the median

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 21

region and with long steep slopes to the margins. Interior with broad,
short sessile spondylium.

Brachial valve gently convex in lateral profile; broadly convex in
anterior profile with the median region somewhat carinated and with
long lateral slopes; umbo narrow and protuberant; median region
inflated gently; fold originating at about the middle, low and sub-
carinate; flanks gently swollen. Interior with short, shallow brachio-
phore plates.

Measurements in mm.

4 Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

ESSN EMER Noe t167A0bense 5. 6.3 ic 8.1 6.8 PLD
Brachial valve,

iS NMS Now TIO740d ye. 5k ? 5.4 7.8 6.0 Dp:

Types.—Holotype: U.S.N.M. No. 116740b; figured paratype:
U.S.N.M. No. 116740d; unfigured paratypes: U.S.N.M. Nos.
1167492, ¢, e.

Horizon and locality—Signal Mountain formation (2-34 feet
above base of Butterly dolomite) in Oklahoma, SW4SW4SWS3 sec. 9,
eos. RI 2. Murray County.

Discussion.—This species is ornamented like G. imbricata from the
Mons formation near Lake Louise, Alberta, but differs in its less-
transverse outline, less-deep pedicle sulcus, and more narrowly
rounded fold on the brachial valve.

IMBRICATIA Cooper, new genus

Shells usually transverse, syntrophoid in profile and outline; an-
terior commissure moderately uniplicate; surface marked by strong
concentric imbrications and fine costellae.

Interior of pedicle valve with small teeth and usually with a thick
spondylium simplex, sessile at the rear but elevated anteriorly and
supported by a short thick median ridge. Vascula media usually
strongly developed.

Brachial valve with short, stout brachiophores and small sockets
defined by small fulcral plates. Brachiophore supporting plates usu-
ally erect to moderately oblique, meeting the floor of the valve;
brachiophore plates usually with thick ridges near the base of the
plate which give them a saucerlike shape; no cardinal process. Ad-
ductor scars as in Tetralobula but usually not with strong callosities.

Genotype.—Imbricatia lamellata Cooper, new species.

Discussion.—This genus is essentially like Tetralobula but differs

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

in having a more elaborate ornamentation which consists of fine
radial costellae interrupted by strong concentric lamellae. Internally
the two genera are similar but differ in details of the brachial in-
terior. The brachiophore supporting plates are alike and were evi-
dently the seat of attachment for diductor muscles. This is clearly
shown by the presence of strong curving ridges near their base which
gives them a saucerlike appearance. The adductor callosities of
Imbricatia are not so strongly developed as those of Tetralobula.

Imbricatia is ornamented like Stichotrophia but differs from that
genus in having brachiophore plates convergent toward the floor of
the valve rather than strongly divergent. In Stichotrophia, further-
more, the brachiophore plates have adductor ridges suggestive of
Imbricatia but they appear high above the floor so that the plate seems
to consist of two elements. Calliglypha Cloud has imbricate ornamen-
tation but in that genus the imbrications and radial lines produce
small spines. Cloud’s genus is readily differentiated internally because
it is like Diaphelasma.

Other species formerly placed in Tetralobula can be assigned to
Imbricatia as follows: Imbricatia coloradoensis (Ulrich and Cooper)
from the Manitou limestone of Colorado; J. imbricata (Ulrich and
Cooper) from the Hastings Creek formation near Philipsburg,
Quebec; and probably Tetralobula dorsosulcata (Ulrich and Cooper)
also from the Manitou limestone of Colorado.

IMBRICATIA LAMELLATA Cooper, new species
Plate 3E, figures 14-19; plate 4D, figures 17-22

Shell of about medium size for the genus, wider than long, roundly
elliptical in outline; hinge wide; posterolateral extremities narrowly
rounded; sides rounded; anterior margin broadly rounded; anterior
commissure gently uniplicate; surface with strong concentric plaits
which are finely costellated ; costellae about eight in one millimeter at
the front margin.

Pedicle valve moderately convex in lateral profile with the maxi-
mum convexity in the anterior half; anterior profile broadly and
moderately convex, umbo and median region swollen ; sulcus shallow,
originating at about the middle; tongue short and narrowly rounded.
Interior with spondylium sessile at rear but strongly elevated ante-
riorly; old shells with pallial marks strongly developed.

Brachial valve moderately convex in lateral profile, broadly and
moderately convex in anterior profile. Fold variable, defined in the
front third only and often appearing as an emargination in the an-

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 23

terior margin. Median region swollen; flanks moderately convex and
with moderate slopes.

Measurements in mm.

3 Brachial Mid- Hinge Thick-
Pedicle valves, Length length width width ness

WESSNEM ae No weti6 7562p aoe. sr 8.7 ? 10.4 9.0 2.4
PU ouN. W UING) TTGZS20) aise a 7.2 ? 8.6 7.6 P42)
Brachial valves,

WESANEME MING ttO75260s).een ? 7.6 10.6 8.4 2.8

WESSNEME wNiossTniG7G20"R sen. ? 8.2 10.9 7.5 3.0

Types—Holotype: U.S.N.M. No. 116752g; figured paratypes:
U.S.N.M. Nos. 116752a, b, 117949a-c; unfigured paratypes:
U.S.N.M. Nos. 116752c-f, h.

Horizon and locality—Middle of Cool Creek formation in Okla-
homa, 100 feet north and 50 feet east of southwest corner of sec. 34,
T.1S., R.1 W., Murray County.

Discussion—This species suggests I. coloradoensis and I. im-
bricata but differs from both of them. From the former it may be
distinguished by its more transverse outline and less sharply folded
brachial valve. It is closer to J. imbricata of Quebec in outline but is
also somewhat more transverse than the species and has a much less
conspicuous and shorter sulcus in the pedicle valve and a usually
more modest development of the brachial fold.

Genus SYNTROPHINA Ulrich, 1928
SYNTROPHINA CAMPBELLI (Walcott)
Plate 3C, figures 4-8
Syntrophia campbelli Waucotr, Smithsonian Misc. Coll., vol. 53, pp. 107, 108,

pl. 10, figs. 9, 9a-c, 1908; U. S. Geol. Surv. Mon. 51, p. 801, fig. 73, 1912.

Syntrophina campbelli (Walcott) Utricn and Cooper, Geol. Soc. Amer. Spec.
Pap. 13, p. 218, pl. 46E, figs. 10-34, 1938.—CLoup, Bull. Mus. Comp. Zool.,
vol. 100, No. 5, p. 462, pl. 2, figs. 10-13, 17, 1948.

Shell small, roundly elliptical in outline, and with the maximum
width at about the middle; sides strongly rounded; anterior com-
missure gently rounded, strongly uniplicate. Surface marked by con-
centric lines of growth.

Pedicle valve moderately convex in lateral profile and broadly and
moderately convex in anterior profile. Umbonal and median region
swollen ; sulcus originating at about the middle, broad and shallow.
Tongue moderately long and narrowly rounded. Flanks bounding
sulcus gently convex and with long steep slopes to the margins.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Spondylium small, short, and supported by a short median septum.
Teeth small, long, narrow.

Brachial valve about equal in depth to the pedicle valve, gently
convex in lateral profile and most convex in the umbonal region;
anterior profile broadly and moderately convex. Umbonal region
swollen; fold originating at about the middle, low and _ broadly
rounded ; flanks bounding fold gently swollen and with steep slopes
to the margins. Interior with broad and short brachiophores sup-
ported by convergent plates that meet the floor of the valve in close
proximity. Sockets defined by small socket plates. Adductor scars
not impressed.

Measurements in mm.

s Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

WS SINE, ING) hile Ozh6s poe auc 7 ? 10.4 5.5 2.5
Brachial valve,

US-NeM. No. 1167402. 5.3... ? 7.5 8.8 4.4 3.0

Types——Figured hypotypes: U.S.N.M. Nos. 116749d-f ; unfigured
paratypes: U.S.N.M. Nos. 116749a-c, g-i.

Horizon and locality—McKenzie Hill formation (360 feet above
the base) in Oklahoma, 950 feet west and 2,050 feet north of the
southwest corner of sec. 26, T. 6 N., R. 14 W., Kindblade Ranch,
Wichita Mountains, Kiowa County.

Discussion—These specimens are unusual in being preserved as
silicified shells. Hitherto this species was known only from chert
impressions. The specimens herein illustrated conform to the details
of the species except for the small transverse plate in the apex of
the brachial valve. This is lacking from the Oklahoma specimens.

Genus CLARKELLA Walcott, 1908
CLARKELLA species
Plate 3B, figure 3

The specimens of this interesting genus reported for the first time
in Oklahoma are unfortunately very fragmentary. Part of a pedicle
valve indicates a species with a deep and wide median sulcus having
a prominent, impressed line running down the middle; the tongue is
long and pointed. The spondylium is strongly elevated on a high
median septum. The brachial valve had a prominent fold but its
prominence did not arise until the anterior part of the shell. The
posterior half is well rounded and without a fold. The interior shows

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 25

the adductor callosities elevated on septa as usual for the genus, and
the diductor ridges just below the notothyrial edge are well developed.

Figured specimen.—U.S.N.M. No. 116750a.

Horizon and locality—Upper Cool Creek formation (300 feet
below the top) in Oklahoma, at the south end of Falls Creek anticline,
2,200 feet north and 2,200 feet west of the southeast corner of sec. 21,
ee Shag e.Catter, County.

Genus DIPARELASMA Ulrich and Cooper, 1936
DIPARELASMA COSTELLATUM Cooper, new species
Plate 3D, figures 9-13

Shell small, subcircular in outline with sides and anterior margins
rounded ; hinge narrow and cardinal extremities obtuse; surface cos-
tellate, three or four costellae to the millimeter at the front margin,
the stronger ones concentrated at the sulcus.

Pedicle valve moderately convex in lateral profile and with the
maximum convexity just anterior to the umbo; anterior profile
strongly convex and with the median region somewhat narrowly
rounded. Umbo narrowly swollen; median region swollen; anterior
regions somewhat flattened ; flanks sloping moderately steeply to the
sides. Delthyrial cavity wide, bounded by strong dental plates.

Brachial valve having less depth than the pedicle valve and gentle
convexity in lateral profile; anterior profile broadly and gently con-
vex ; umbo swollen and median and anterior regions moderately in-
flated; sulcus originating on the umbo, narrow and shallow, extend-
ing to the anterior margin. Flanks gently swollen; lateral slopes
gentle.

Measurements in mm.

y Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

MESIN ME INO: | EOZSAR ie «aja, 010 « 6.3 ? 7.6 5.0 23
Brachial valve,

WO SeNeViE SNOyoniG75Abeaseseers ? 6.0 7.6 4.9 1.2

Types.—Holotype: U.S.N.M. No. 116754a; figured paratypes:
U.S.N.M. Nos. 116754b, c.

Horizon and locality—West Spring Creek formation (419 feet
below the top) in Oklahoma, 1,400 feet east and 1,700 feet north of
the southwest corner of sec. 6, T. 3 S., R. 4 E., Johnston County.

Discussion.—This species is characterized by small size, moderate
convexity, and fairly strong costellae. The species is most like D.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IL7,

typicum of described species but is smaller, less convex, and more
strongly costellate particularly in the median region.

DIPARELASMA FASCICULATUM Cooper, new species
Plate 4B, figures 8-12

Shell of about usual size for the genus, subcircular in outline ; sides
and anterior margin rounded ; hinge narrower than the greatest shell
width which is at the middle. Anterior commissure slightly sulcate.
Surface costellate, fasciculate in bundles of two to four costellae.

Pedicle valve moderately convex in lateral profile and with the
maximum convexity just anterior to the umbo, the remainder of the
valve more gently convex; anterior profile narrowly convex in the
median region and with steep lateral slopes. Umbonal and median
regions swollen, the swelling extending anteriorly to the front mar-
gin to form a faint fold. Interior with dental plates well developed.

Brachial valve gently convex in lateral profile and with the maxi-
mum convexity just anterior to the umbo; anterior profile gently and
broadly convex but with the median region slightly depressed. Umbo
moderately swollen; sulcus originating just anterior to the umbo,
shallow, widening anteriorly but remaining shallow throughout, and
occupying about half the width at the front margin. Tongue short,
broadly rounded; flanks bounding sulcus slightly swollen and with
gentle slopes to the margins. Interior with short and delicate brachio-
phore plates and short, low median ridge.

Measurements in mum.

; Brachial Mid- Hinge Thick-
Pedicle valve, Length length width width ness

WESSNEMe Nigh Ta 7O4Sarere tears 6.7 ? 6.9 4.9 2.3
Brachial valve,

USS. NM UNG. TEZO48be ee eects ? 6.8 8.2 5.7 1.6

Types.—Holotype: U.S.N.M. No. 117948b; paratype: U.S.N.M.
No. 117948a.

Horizon and locality—Upper Kindblade formation (about 250 feet
below the top) in Oklahoma, 700 feet east and 1,200 feet south of
the northwest corner of sec. 5, T. 3 S., R. 4 E., Johnston County.

Discussion.—This species is characterized by its fasciculate cos-
tellae and the short brachiophore plates. It differs from D. costel-
latum in having the finer costellae concentrated in the sulcus and in
having much more delicate cardinalia. It differs from all other de-
scribed species in its fasciculate ornamentation.

NO. 14 BRACHIOPODS FROM OKLAHOMA—COOPER 27

POLYTOECHIA SUBCIRCULARIS Cooper, new species
Plate 4C, figures 13-16

Shell wider than long, circular in outline and with the hinge nar-
rower than the greatest shell width which is at the middle; sides
gently rounded ; anterior commissure rectimarginate ; surface multi-
costellate, seven costellae in one millimeter at the anterior margin.

Pedicle valve fairly evenly convex in lateral profile and with the
maximum convexity at about the middle; anterior profile strongly
convex; beak small, narrowly swollen; umbo somewhat narrowly
swollen; median region inflated and most strongly convex near the
center; sides steeply sloping to the margins; anterior slope long and
less steep than the lateral slopes. Interarea long, curved, apsacline.
Interior with strong dental plates, erect and subparallel; muscular
area slightly elevated above the floor of the valve; diductor scars
elongate ; adductor region marked by a long narrow groove bounded
by two plates convergent with the median septum and extending an-
terior to the delthyrial cavity. Dental plates extended as slightly diver-
gent ridges anteriorly to the anterior end of the median septum. Pal-
lial trunks not clearly defined anterior to the ends of the extensions of
the dental plates.

Brachial valve with evenly and moderately convex lateral profile ;
anterior profile broadly and moderately convex ; valve broadly ellipti-
cal in outline; umbo and median regions strongly swollen; lateral
slopes steep and moderately long; anterior slope long, gently rounded
and less steep than the lateral slopes. Interior with short convergent
brachiophore plates, small cardinal process, short median ridge but
lightly impressed pallial and muscular marks.

Measurements in mm.

$ Brachial Mid- Hinge Thick-
Pedicle valves, Length length width width ness
IU SeNe Me Nios 17052a\..4-1)--14- 8.2 ? 9.2 6.7 3:2
WESIN aVisINow tI7O52C>.4- li ? Gi 10.9 7.8 2.4

Types.—Holotype: U.S.N.M. No. 117952a; figured paratypes:
U.S.N.M. Nos. 117952b, c; unfigured paratypes: U.S.N.M. Nos.
117952d-f.

Horizon and locality—West Spring Creek formation (598 feet
above the base) in Oklahoma, on Joins Ranch, 750 feet west and
1,550 feet south of the northeast corner of sec. 9, T. 2 S., R. 1 W.,
Murray County.

Discussion—This species is characterized by its nearly circular

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. LL

pedicle valve, somewhat narrowly elliptical brachial valve, very fine
costellae, and the low triseptate spondylium in the pedicle valve. This
species suggests P. alabamensis and P. fillistriata in its fine costellae
but differs from the former in being smaller, having a longer interarea
which is less curved and a much lower spondylium. The Oklahoma
species differs from P. fillistriata in having somewhat finer ornamen-
tation and an extremely low spondylium, that of the Newala shell
being strongly elevated.

SELECTED REFERENCES
Gxoup, PB:
1948. Brachiopods from the Lower Ordovician of Texas. Bull. Mus. Comp.
Zool., vol. 100, No. 5, pp. 451-475, pls. 1-4.
Cooper, G. A.
1948. A new genus of Brachiopoda from the Longview limestone of Vir-
ginia. Bull. Mus. Comp. Zool., No. 6, pp. 473-474, pl. 4.
DECKER, CE.
1939. Two lower Paleozoic groups, Arbuckle and Wichita Mountains,
Oklahoma. Bull. Geol. Soc. Amer., vol. 50, pp. 1311-1322.
Unricy, E. O., and Cooper, G. A.
1938. Ozarkian and Canadian Brachiopoda. Geol. Soc. Amer. Spec. Pap.
13, Ppp. i-vili, 1-323, pls. 1-58.
Watcort, C. D.
1912, Cambrian Brachiopoda. U. S. Geol. Surv. Mon. 51, pp. 725-810, pls.
85-104.

EXPLANATION OF PLATES

PLATE I

A. Buillingsella rectangulata Cooper, new species.............ceceeccecces

1, Brachial view of a complete specimen showing pseudodel-
tidium, X 1; paratype, U.S.N.M. No. 116722b. 2-6, 8, Pos-
terior tilted, pedicle, posterior, anterior, side, and brachial
views, respectively, of the holotype, U.S.N.M. No. 116722a,
<2. 7, The holotype, XI. 9, I0, Exterior and interior
of the brachial valve, X 2, showing brachiophores; para-
type, U.S.N.M. No. 116722k. 11, Interior of the pedicle
valve showing teeth, vascula media, and thickening anterior
to muscle scar, X 2; paratype, U.S.N.M. No. 116722e. 12,
Another brachial interior showing notothyrial platform
and median ridge, X 2; paratype, U.S.N.M. No. 116722i.
13, 14, Pedicle valve showing teeth and pallial marks, and
same tilted to show callus supporting teeth, 2; para-
type, U.S.N.M. No. 116722d. 15, Another pedicle interior
showing teeth; paratype, U.S.N.M. No. 116722h.

Signal Mountain formation (1-foot bed of limestone 100 feet
above base), 1,700 feet east and 1,300 feet north of south-
west corner of sec. 21, T. 1 S., R. 1 W., Murray County,
Okla.

Be Cymbithgns ham Cooper, New SPeCieS:. i... oe. cc eee ne cece eee sens ee

16, Imperfect specimen showing both valves in contact and with
missing parts restored, 2; holotype, U.S.N.M. No.
116724d. 17, Exterior of a nearly complete pedicle valve,
2; paratype, U.S.N.M. No. 116724a. 18, Exterior of
another pedicle valve showing form, 2; paratype,
U.S.N.M. No. 116724b. 19, Exterior of a third pedicle
valve, X 1; paratype, U.S.N.M. No. 116724e. 20, Interior
of the brachial valve showing reduced brachiophores, x 2;
paratype, U.S.N.M. No. 116724f. 21, Interior of the pedicle
valve showing teeth and pseudodeltidium, 2; paratype,
U.S.N.M. No. 116724c.

Signal Mountain formation (limestone 1 foot thick 126 feet
above base), 1,700 feet west and 600 feet south of the
northeast corner of sec. 28, T. 1 S., R. 1 W., Murray
County, Okla.

C. Finkelnburgia crassicostellata Cooper, new species............00---00.

22, Exterior of the pedicle valve showing costellae, & 1; holo-
type, U.S.N.M. No. 116726a. 23, Same, X 2, showing
coarse and fine costellae. 24, Interior of the pedicle valve,
X 2, showing broad pseudospondylium; paratype, U.S.N.M.
No. 116726b. 25, 26, Exterior and interior, respectively, of
the brachial valve showing ornamentation and adductor
callosities, X 2; paratype, U.S.N.M. No. 116726c.

D. Mesononuia magna Cooper, new species
27, Exterior of the pedicle valve showing the ornamentation,

A. Fasciculina fasciculata Cooper,,new genus and species
I, 2, Exterior and interior of the pedicle valve showing fascic-

B. Apheoorthis platys Cooper, new species
8, 9, Exterior of the pedicle valve, X I and X 2, respectively,

C. Glyptotrophia rotunda Cooper, new species
14, Exterior of the brachial valve showing fold and ornamenta-

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. ui,

Kindblade formation, in center NE{SW34 sec. 29, T. 1 S.,

R. 1 W., Murray County, Okla.

<1; holotype, U.S.N.M. No. 116694a. 28, 29, Exterior
and interior, respectively, of the holotype showing details
of the ornamentation, the teeth, and muscle region, X 2.
30, Brachial interior, X 2, showing cardinal process; para-
type, U.S.N.M. No. 116604c. 31, Exterior of a nearly per-
fect pedicle valve, X 1; paratype, U.S.N.M. No. 116694b.
32, 33, Exterior and interior, respectively, of the preceding,
2, showing ornamentation and brachiophores and their
supporting plates. 34, Interior of a second pedicle valve
showing vascula media, 2; paratype, U.S.N.M. No.
116694d.

Fort Sill formation (179 feet above the base), 1,000 feet south

of the north quarter corner of sec. 20, T. 1 S., R. 1 E,,
Murray County, Okla.

PLATE 2

ulate costellae and cardinalia, 2; holotype, U.S.N.M.
No. 116738e. 3, Exterior of the pedicle valve, & 2, show-
ing median fascicles; paratype, U.S.N.M. No. 116738b.
4, Interior of another pedicle valve showing pseudospondyl-
ium, <2; paratype, U.S.N.M. No. 116738c. 5, 6, Same
views as in figures I and 3 showing brachial and pedicle
exterior, X I. 7, Interior of a brachial valve showing cardi-
nalia; paratype, U.S.N.M. No. 116738d.

Signal Mountain formation (90 feet above the base),

SESSWiNE4 sec. 16, T. 1 S., R. 1 W., 1,200 feet west of
east quarter corner, Murray County, Okla.

showing the ornamentation; holotype, U.S.N.M. No.
116736a. 10, Interior of the holotype showing pseudo-
spondylium and vascula media, X 2. II, 12, Exterior of
the brachial valve, X1 and 2; paratype, U.S.N.M.
No. 116736b. 13, Interior of the same brachial valve as the
preceding showing the orthoid cardinalia, x 2.

Signal Mountain formation (50 feet below the top), 2,100 feet

north and 100 feet west of the southeast corner of sec. 26,
T..1,S8,,,R., 0 W., -Murray, County, Okla:

tion, X 2; paratype, U.S.N.M. No. 116740d. 15, Exterior
of the pedicle valve showing short and shallow sulcus, X 2;
holotype, U.S.N.M. No. 116740b.

Page

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER 31

Page
Signal Mountain formation (2-34 feet above base of "Butterly
dolomite), SW4SWiSW3 sec. 9, T. 2 S., R. 2 E., Murray
County, Okla.
D. Finkelnburgia delicatula Cooper, new species............ceeecccecscees 14
16, Exterior of the brachial valve, X 1; paratype, U.S.N.M.
No. 116734f. 17, 18, Interior and exterior, respectively, of
the same specimen showing details of the adductor cal-
losities, the cardinalia, and the ornamentation, 2. Io, In-
terior of another brachial valve, & 2, showing adductor
callosities occuping center of valve; paratype, U.S.N.M.
No. 116734g. 20, 21, Views of the exterior of the pedicle
valve, X1 and 2, respectively; holotype, U.S.N.M.
No. 116734a.
Upper Cool Creek formation, 1 mile southeast of the windmill,
center of sec. 5, T. 2 S., R. 1 E., Murray County, Okla.
E. Finkelnburgia biconvexa Cooper, new SpeCieS...........ceecceeeecnces 12
22, 23, Views of the exterior of the pedicle valve, 1 and
< 2, respectively, showing details of the ornamentation;
holotype, U.S.N.M. No. 116733a. 24, Interior of the holo-
type showing the low pseudospondylium, X 2. 25, Incom-
plete pedicle valve showing pseudospondylium, XX 2; para-
type, U.S.N.M. No. 116733c. 26, Interior of an imperfect
brachial valve showing cardinalia, X 2; paratype, U.S.N.M.
No. 116733e. 27, 28, Interior and exterior of a brachial
valve, 2, showing cardinalia and details of the orna-
mentation; paratype, U.S.N.M. No. 1167336.
Signal Mountain formation (218 feet above the base), 500 feet
west and 750 feet south of the northeast corner of sec. 28,
T. 1 S.. R. 1 W., Murray County, Okla.
F. Finkelnburgia auriculata Cooper, new SpeCi€S...........eeeeeecceeeees 10
29-33, Brachial, anterior, posterior, side, and pedicle views,
respectively, of the holotype, X 2; U.S.N.M. No. 11673!a.
34, 38, Exterior and interior views of a pedicle valve,
X 2, showing low pseudospondylium; paratype, U.S.N.M.
No. 116731e. 35, Interior view of another pedicle valve
showing auriculation of ears, <2; paratype, U.S.N.M.
No. 116731f. 36, 37, Interior views of two brachial valves,
X 2, showing cardinalia and adductor callosities ; paratypes,
U.S.N.M. Nos. 1167314, c.
Signal Mountain formation (6-inch bed of limestone 150 feet
above the base), 300 feet west and 1,150 feet south of the
northeast corner of sec. 28, T. 1 S., R. 1 W., Murray
County, Okla.
G. Finkelnburgia extensa Cooper, new SpeCieS....... eee cece ceceeeee ces 15
30, 40, Interior view of the same pedicle valve tilted and in
normal view to show the teeth and pseudospondylium, X 2;
holotype, U.S.N.M. No. 116728b. 41, 42, Interior view of
a brachial valve, tilted and in normal view to show cardi-
nalia, X 2; paratype, U.S.N.M. No, 116728a. 43, Interior

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

)

of a pedicle valve showing deep pallial channels, X 2;
paratype, U.S.N.M. No. 116728e. 44, Interior of a brachial
valve having long auriculations and showing sockets and
brachiophores, X 2; paratype, U.S.N.M. No. 116728f. 45,
46, Exterior and interior of pedicle valve showing large
thick teeth, pseudospondylium, vascula media, and auricula-
tions, X 2; paratype, U.S.N.M. No. 116728j. 47, Exterior
of an exceptionally large and extended pedicle valve, « I;
paratype, U.S.N.M. No. 116728g. 48, 49, Same as preced-
ing, X 2, to show details of ornamentation and some of the
pallial marks. 50, 51, Views of the exterior of a brachial
valve, X 1 and & 2, respectively, showing shallow sulcus ;
paratype, U.S.N.M. No. 116728d.

Signal Mountain formation (90 feet above the base),
SESSWiNE? sec. 16, T. 1 S., R. 1 W., 1,200 feet west
of the east quarter corner, Murray County, Okla.

PLATE 3

INS JEollorqossAowiitor Cin J Ate (OEMS) sacccndeascondsducaonunecsasomed

1, Exterior of an imperfect pedicle valve, 1; figured speci-
men, U.S.N.M. No. 116751a. 2, Interior of the brachial
valve showing the cardinalia, 2; figured specimen,
U.S.N.M. No. 116751b.

Signal Mountain formation (63 feet below the top), 1,500 feet
west and 1,900 feet north of the southeast corner of sec. 6,
T.2S., R. 2 E, Murray County, Okla.

By ClarRella, (Spectes) a cistsce fea sy cts lars, aero des ieysaey iat ess Spore beiaciot= kets iste papsk one

3, Interior of an imperfect brachial valve showing cardinalia,
x 2; figured specimen, U.S.N.M. No. 116750a.

Upper Cool Creek formation (300 feet below the top), south
end of Falls Creek anticline, 2,200 feet north and 2,200 feet
west of the southeast corner of sec. 21, T. 2 S., R. 2 E,,
Carter County, Okla.

Caysyntrophinagcampvellin (NVialcott) hneetac one cere ecco

4, Interior view of a brachial valve tilted to show the cardi-
nalia, X 2; hypotype, U.S.N.M. No. 116749f. 5, Exterior
view of preceding, & 2. 6, Interior view of a pedicle valve
showing pseudospondylium and vascula media, 2; hypo-
type, U.S.N.M. No. 116749e. 7, 8, Interior and exterior
views of another pedicle valve showing vascula media and
exterior sulcus; hypotype, U.S.N.M. No. 116740d.

McKenzie Hill formation (360 feet above the base), 950 feet
west and 2,050 feet north of the southwest corner of sec. 26,
T. 6 N., R. 14 W., Kindblade Ranch, Wichita Mountains,
Kiowa County, Okla.

D. Diparelasma costellatum Cooper, new species.......-..eeeeeeeeeeees

9, 10, Exterior views of the pedicle and brachial valves, re-
spectively, 1; holotype, U.S.N.M. No. 116754a, and
paratype, U.S.N.M. No. 116754b. 11, 12, Exterior views of

Page

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER

the same two specimens, 2, showing details of the orna-
mentation. 13, Interior view of a brachial valve showing
the cardinalia, X 2; paratype, U.S.N.M. No. 116754c.

West Spring Creek formation (419 feet below the top), 1,400
feet east and 1,700 feet north of the southwest corner of
sec. 6, T. 3 S., R. 4 E., Johnston County, Okla.

E. Imbricatia lamellata Cooper, new genus and species...............e00-

14, 15, Exterior views of the pedicle and brachial valves, « 1;
paratype, U.S.N.M. No. 116752b, and holotype, U.S.N.M.
No. 116752g. 16, Exterior view of the preceding pedicle
valve, X 2, showing the ornamentation. 17, Interior view
of an old pedicle valve showing the pseudospondylium,
vascula media, and other pallial marks, * 2; paratype,
U.S.N.M. No. 116752a. 18, 190, Exterior and interior of
the holotype, 2, showing details of the ornamentation
and the low adductor callosities.

Middle of the Cool Creek formation, 100 feet north and 50 feet
east of the southwest corner of sec. 34, T. 1 S., R. 1 W.,
Murray County, Okla.

iRNmeeielnour gran ogesa (Gloudiin sats cere isie oc <5 cleierajarclalatets 5 <1atehajersier isis scree

20, 21, Exterior views of the pedicle and brachial valves, re-
spectively, X 1; hypotypes U.S.N.M. Nos. 116745b, d. 22,
23, Interior and exterior views of the same hypotype show-
ing the pseudospondylium and vascula media, X 2. 24, 25,
Interior and exterior views, respectively, of the brachial
valve showing cardinalia and adductor callosities, XX 2;
hypotype, U.S.N.M. No. 116745d. 26, 27, Interior views
of two brachial valves showing cardinalia, adductor cal-
losities, and pallial markings; hypotypes, U.S.N.M. Nos.
116745¢, c.

McKenzie Hill formation (35 feet below the top), 2,200 feet
east and 2,500 feet north of the southwest corner of sec.
2, T.2S., R.1 E., Murray County, Okla.

G. Finkelnburgia ci. F. bellatula Ulrich and Cooper......................

28, 29, Brachial and pedicle exteriors, respectively, 1; hypo-
types, U.S.N.M. Nos. 116747b, a. 30, Interior of the pre-
ceding brachial valve showing the short adductor field and
cardinalia, 2. 31, 32, The preceding pedicle valve in
interior and exterior views to show pseudospondylium and
details of the ornamentation, x 2. ‘

McKenzie Hill formation (10 feet below the top), 2,200 feet
north and 1,250 feet west of the southeast corner of sec. 7,
T. 1 S., R. 1 E., Murray County, Okla.

H. Finkelnburgia arbucklensis Cooper, new SPECIES... .....ccce cece cecees

33, 34, Views of a pedicle valve, X 1 and X 2, respectively,
showing the ornamentation; holotype, U.S.N.M. No.
116741b. 35, Exterior of a brachial valve, X 1; paratype,
U.S.N.M. No. 116741d. 36, 37, Exterior and interior views
of the same brachial valve as the preceding, X 2, showing

Page

a4 F SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

cardinalia, adductor field, and pallial markings. 38, Interior
view of a pedicle valve showing the stout and low pseudo-
spondylium and the deep vascula media, 2; paratype,
U.S.N.M. No. 116741a. 309, Interior view of a brachial
valve, < 2, showing adductor field and cardinalia; para-
type, U.S.N.M. No. 11674Ic.

McKenzie Hill formation (237 feet above the base), 1,500 feet
east and 2,000 feet south of the northwest corner of sec.
28, T. 1 S., R. 1 W., Murray County, Okla.

I. Finkelnburgia subquadrata Cooper, new SpeCieS..........2.eeeeeeeee

40, Exterior of a pedicle valve, 1; holotype, U.S.N.M.
No. 116743a. 41, 42, Interior and exterior views of the
holotype, 2, showing short, thick pseudospondylium and
deep, widely divergent vascula media. 43, Exterior of a
brachial valve, X1; paratype, U.S.N.M., No. 116743b.
44, 45, Exterior and interior of the same brachial valve
as the preceding showing cardinalia and adductor callosities.

Cool Creek formation, 900 feet west and 1,450 feet north of the
southeast corner of sec. 3, T. 2 S., R. 1 W., Murray
County, Okla.

PLATE 4

AO hetralobulavterana Ulrich andi Gooperseenen sete eee

I, 2, Pedicle and brachial views, respectively, 1, of adult
specimens ; figured specimens, U.S.N.M. Nos. 117950e, f. 3,
Exterior of the pedicle valve showing the fine costellae,
X 2; figured specimen, U.S.N.M. No. 117950a. 4, Exterior
of the brachial valve, 2; figured specimen, U.S.N.M.
No. 117950f. 5, Exterior of a young brachial valve, X 3,
showing ornamentation; figured specimen, U.S.N.M. No.
117950b. 6, Interior of the brachial valve showing the ad-
ductor callosities, <3; figured specimen, U.S.N.M. No.
117950d. 7, Interior of the pedicle valve, & 3, showing the
strong pallial trunks and the spondylium; figured specimen,
U.S.N.M. No. 117950c.

McKenzie Hill formation (about 250 feet above the base),
2,100 feet west and 1,500 feet north of the southeast cor-
ner of sec. 36, T. 1 S., R. 1 E., Murray County, Okla.

B. Diparelasma fasciculatum Cooper, new SpeCicS....:....0..eeeeeccees

8, 9, Exterior views of the pedicle and brachial valves, respec-
tively, X 1; paratype, U.S.N.M. No. 117948a, and holotype,
U.S.N.M. No. 117948b. 10, 12, Same views of same speci-
mens, X 2. 11, Interior of the preceding brachial valve, X 2,
showing short brachiophore.

Upper Kindblade formation (250 feet below the top), 700 feet
east and 1,200 feet south of the northwest corner of sec. 5,
1.3 S., R., 4). Johnston Gountys Okla.

Page

NO. I4 BRACHIOPODS FROM OKLAHOMA—COOPER

GC) Polvtoechiasubcircularis Cooper; New speciesa....... ss seee6ssceeesens

13, Exterior of the brachial valve, < 3, showing the fine cos-
tellae; paratype, U.S.N.M. No. 117952c. 14, Interior of
the brachial valve showing the convergent brachiophore
plates like those of Tritoechia, X 3; paratype, U.S.N.M.
No. 117952b. 15, 16, Normal view and view of the valve
tilted, respectively, to show the spondylium supported by
the dental plates and a median septum, X 3; holotype,
U.S.N.M. No. 117952.

West Spring Creek formation (598 feet above the base), on
the Joins Ranch, 750 feet west and 1,550 feet south of the
northeast corner of sec. 9, T. 2 S., R. 1 W., Murray
County, Okla.

D. Imbricatia lamellata Cooper, new genus and species.................05

17, Exterior of the pedicle valve showing the imbrications,
2; paratype, U.S.N.M. No. 117949a. 22, Interior of the
same pedicle valve showing the spondylium, x 2. 18, Io,
Exterior and side views of the brachial valve showing im-
bricate ornamentation, 2; paratype, U.S.N.M. No.
117949c. 20, 21, Tilted and normal views of another bra-
chial valve showing cardinalia, X 2; paratype, U.S.N.M.
No. 117940b.

Cool Creek formation (300 feet below the top), center NE}
sec. 6, T. 2 S., R. 1 E., Murray County, Okla.

Ex eeiectowophia latucosta @ooper, BREW. SPECIES... Sockets osc ees

23, 24, Exterior views of the pedicle and brachial valves, re-
spectively, X 1; paratypes, U.S.N.M. Nos. 117951d, c. 25,
The same pedicle valve, X 2, showing sparse costellae. 26,
Exterior of the brachial valve showing low fold and distant
costellae, X 2; holotype, U.S.N.M. No. 117951g. 27, An-
other brachial valve showing the same features, X 2; para-
type, U.S.N.M. No. 117951e. 28, 29, Exterior and side
views of the brachial valve showing the low fold and
distant costellae, 3 and XX 2, respectively; paratype,
U.S.N.M. No. 117951f. 30, 31, Interior of the brachial
valve showing the cardinalia, normal view, and the same
valve tilted, respectively, 2; paratype, U.S.N.M. No.
117951b. 32, 33, Tilted valve and normal view of the
pedicle interior, respectively, showing tapering spondylium
and adductor track, & 3; paratype, U.S.N.M. No. 117951a.
34, 35, Normal interior view and the same valve tilted,
respectively, to show the converging brachiophore plates,
<3; paratype, U.S.N.M. No. 117951b.

Fort Sill formation (150 feet above the base), 2,500 feet east
and 2,200 feet north of the southwest corner of sec. 26,
T.2S., R. 4 E., Johnston County, Okla.

eae tc ee Cer a le ee Le Wie JIN
A fen? SF ied i | a ive ya 1
’ 7 t ; nt Vine AY iy havo a

fail! a ieat
i.e iis
; aes
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ry
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ay ;
bey
' , pe eal ‘if
4 RAgGes}
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, Py OM i Hat tah Ahh a

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 14, PL. 1

BRACHIOPODS FROM THE ARBUCKLE GROUP, OKLAHOMA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 14, PL. 2

eg? | Nae

BRACHIOPODS FROM THE ARBUCKLE GROUP, OKLAHOMA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 14, PL. 3

BRACHIOPODS FROM THE ARBUCKLE GROUP, OKLAHOMA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 14, PL. 4

BRACHIOPODS FROM THE ARBUCKLE GROUP, OKLAHOMA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 15

soy hae DB. and Mary Waux Walcott
| Research Fund

_ THE FORAMINIFERAL GENUS
TRIPLASIA REUSS, 1854

(WitH 8 PratTEs)

BY
ALFRED R. LOEBLICH, JR.
AND

HELEN TAPPAN
U. S. National Museum

eee eeces,

Seecee0ee?

( Pustication 4094)

ees! CITY OF WASHINGTON
"PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 9, 1952

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 15

Charles D. and Mary Waux Walcott
Research Fund

foe FORAMINIPERAL GENUS
TRIPLASIA REUSS, 1854

(WitH 8 PLaTEs)

BY
ALFRED R. LOEBLICH, JR.
AND

HELEN TAPPAN
U. S. National Museum

5), Net
THSOM SS,
4 fiapeas Cone

NCHINGTO Lp

@ecee0ee®

(Pus.ication 4094)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 9, 1952

The Lord Baltimore Press

BALTIMORE, MD., U. 8. A.

Charles D. and Mary Waux Walcott Research Fund

THE FORAMINIFERAL GENUS TRIPLASIA
RUSS 5.1854
By ALFRED R. LOEBLICH, JR. ann HELEN TAPPAN

U. S. National Museum
(With Eicut PLates)

INTRODUCTION

Thirty-seven species of Triplasia, including nine new species,
are described herein, and most of these are figured, from strata
ranging in age from Lower Jurassic to Recent. An examination
of abundant material shows a definite gradation within many of
the species from forms with a triangular section to 4-sided specimens.
Furthermore, the early planispiral coil is extremely variable in size
and prominence, and it may be lacking entirely. These gradations,
which are more fully discussed under the species concerned, are
also shown in the illustrations and have led the writers not only
to agree with Bartenstein and Brand (1951) in suppressing the genus
Frankeina as a synonym of Triplasia but also similarly to suppress the
genera based on quadrate forms, Centenarina Majzon and Tetraplasia
Bartenstein and Brand.

Triplasia Reuss was defined in 1854, the genotype species being
Triplasia murchisoni. Ruess’s original description stated (transla-
tion from German) :

Test free, regular, straight, inverted-ovate or somewhat elongate in outline,
3-sided. Chambers placed in a straight line upon each other, identical, some-
what equitant, not separated by constrictions. The final chamber is produced
into a short central neck, which carries the round aperture.

Differs from Nodosaria in the 3-sided form, with identical unconstricted,
somewhat equitant chambers, and the rough, not glassy, shell wall, and from
Orthocerina by the equitant chambers and the central prolongation of the final
chamber.

Before additional species were described, Reuss defined the genus
Rhabdogonium (1860), placing Triplasia in it as a synonym, and stat-
ing in part (translation from German) :

The most conspicuous character of Rhabdogonium is the presence of sharper
longitudinal angles on the straight test. The four first-known forms possessed

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 15

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

only three such angles, were also triangular in section, wherefore I gave the
genus the name Tyriplasia. Here belong: Rh. murchisoni m., from the Gosau-
schicten from the Austrian Saltz domain, Rh. globuliferwm, roemeri, and
anomalum m., which I forthwith describe from the Cretaceous of Westphalia
and finally Rh. acutangulum m., an as yet unpublished species that I discovered
in the Hils of Berklingen.

Later, however, I discovered 4-sided species that, except for the tetragonal
cross section, agree fully with the aforementioned in their remaining character-
istics, as Rh. strombecki and mertensi m., both also occurring in the Hils of
Berklingen and not as yet published. For these the earlier name Triplasia obvi-
ously cannot now be employed, and I therefore am compelled to substitute a
new one, Rhabdogonium. In this way the angled rhabdoidal form of the test
would be clearly signified.

Reuss therefore renamed his genus when he discovered quadrate
specimens which he thought to be congeneric. This cannot be done,
for the Rules of Nomenclature require that the first valid name be
kept, regardless of its suitability, and Triplasia must therefore be
retained for the genus of Foraminifera to which belongs T. murchisont,
the genotype species.

Although the first species he mentioned in connection with the
name Rhabdogonium was R. murchisoni (the genotype species of
Triplasia), Reuss also included in his discussion some triangular
calcareous species (R. globuliferum, R. anomalum, and R. acutangu-
lum), all of which are now considered to belong to the genus Tristix
Macfadyen. Furthermore, and ironically, the two quadrate species,
whose discovery led Reuss to rename his genus, also were calcareous
and both are now placed in Tristix, as intraspecific variations of
triangular species.

Because Reuss had so quickly renamed his genus, he was followed
by many workers, and the arenaceous triangular and quadrate species
were for nearly a century referred either to Rhabdogonium or Haplo-
phragmium. The first species referred to Triplasia (other than the
genotype species) was T. manderstjent Costa, 1894. This species,
however, was a calcareous, coiled form, and undoubtedly a Sara-
cenaria. The next species to be referred to the genus were T. reussi
Cushman, also a calcareous form and probably a Trifarina, and T.
wilcoxensis Cushman and Ponton, 1932, T. temirica Dain, 1934, and
T. somaliensis Macfadyen, 1935, which are all calcareous, uniserial,
triangular lagenids, now referable to the genus Tristix.

Agglutinated and calcareous species were included in the same
genus by many of the earlier workers, including Reuss, who had not
considered wall characters to be diagnostic. Apparently the later

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 3

workers noted the calcareous species referred to Triplasia and Rhab-
dogonium by their author and ignored the arenaceous character of
the type species.

As late as 1932 Cushman considered Triplasia to be a calcareous
genus, and Cushman and Ponton described T. wilcoxensis in 1932,
stating, “It may be referred to Reuss’ genus Triplasia which was
afterward renamed by him Rhabdogonium because he later found
specimens which had more than three sides . . . It is apparently uni-
serial throughout and belongs with the Lagenidae.” This species now
is placed in Tristix, as is mentioned above.

The genus Frankeina was described by Cushman and Alexander
(1929, p. 61) for arenaceous forms with an early coil and triangular
uniserial later portion, and was placed in the Lituolidae with the
genotype species F'. goodlandensis, from the Lower Cretaceous (Al-
bian) Goodland formation of Texas (erroneously stated in original
publication to be from the “Upper Cretaceous, Upper Middle Good-
land formation”). Cushman and Alexander also called attention to
the peculiar living South Pacific species, Verneuilina variabilis Brady,
stating that it might possibly be a living example of Frankeina, but
further study was necessary to prove this point. It was also pointed
out that some Upper Cretacecous triangular forms referred to Clavu-
lina might not have an early triserial portion and had their origin in
Frankeina, by acceleration of development and resultant loss of the
early coil.

After a trip to Germany and Austria in 1932 Cushman placed Tri-
plasia (and Rhabdogonium as a synonym) in the Lituolidae (Foram-
inifera, 2d ed., 1933), describing it as “triangular or quadrangular
in section, early stages in the microspheric form planispirally coiled,
in the megalospheric form uniserial throughout; ... wall rather
coarsely arenaceous ...” He further stated that “Study of type material
in Vienna showed that this species is derived from Frankeina, by the
loss of the coiled stage in the megalospheric form.” Later, however,
Cushman (1946, p. 26) called attention to this reduction of the coil
in the species Frankeina taylorensis Cushman and Waters, stating
that little more acceleration would eliminate the coil entirely and that
it would be difficult to distinguish these forms from Pseudoclavulina
unless their early portions were sectioned. However, if Triplasia
represented those species in which the coil is lost in the megalospheric
generation, as stated by Cushman in his text, then this species and
similar ones should have been referred to Triplasia. Had Cushman’s

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 1L7

reexamination of the European material occurred before the descrip-
tion of the numerous American species of Frankeina, it seems very
probable that the latter would have been originally placed in Triplasia.
However, as the genotype species represented opposite ends of the
gradational series of species, Frankeina goodlandensis with an ex-
ceptionally well-developed coil, and Triplasia murchisoni lacking or
with a very poorly developed coil, both genera were recognized by
Cushman in his texts of 1933, 1940, and 1948.

In practice, however, American workers have apparently referred
all species to Frankeina, and European workers have recently placed
their species in Triplasia, a duplication in nomenclature which obvi-
ously should be discontinued.

While this paper was being prepared with the intention of sup-
pressing Frankeina, Centenarina, and Tetraplasia all as synonyms of
Triplasia, the publication of Bartenstein and Brand (1951) appeared,
in which for the first time Frankeina was suppressed as a synonym
of Triplasia, although the quadrate forms were separated as species
of the genus Tetraplasia.

Bartenstein and Brand stated (1951, p. 273, translation from
German) :

In the foreign literature Tviplasia-like tests from the Lower and Upper Creta-
ceous have been placed as a rule in the genus Frankeina Cushman and Alexander,
1929... . [quoting Cushman’s statement that Triplasia is derived from Frank-
eina by the loss of the coil in the megalospheric form]. . . . This distinction we
do not consider sufficient as both genera agree fully in all other characters of
the test. We therefore believe that Frankeina must be suppressed as a synonym
of Triplasia.

Reuss considered the number of angles in the test to be of specific
importance, but not generic. Two later workers, Beissel and Brady,
described triangular and quadrate specimens as conspecific as well. As
the quadrate forms are much less common, little attention was paid
to them by most writers, although a few quadrate species and varie-
ties were described. In the past four years two generic names were
proposed independently for these quadrate species (Centenarina
Majzon, 1948, and Tetraplasia Bartenstein and Brand, 1949) and
in both publications the quadrate specimens figured by either Beissel
or Brady were mentioned.

The present writers have examined numerous species of these
rhomboid lituolids, and in every instance, whenever a large suite of
specimens was available, each species was found to show gradations
from triangular to quadrate sections. This is also true for all geologic

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 5

horizons, the earliest known species from the Lias, T. kingakensis,
new species, showing some flattened, some triangular, and some quad-
rate specimens, and the youngest species, the Recent T. variabilis
(Brady), showing a similar gradation. The genotype species of
Frankeina, F. goodlandensis, shows the same gradation, and the type
species of Tetraplasia, T. georgsdorfensis, is conspecific with triangular
forms from the same strata. The type species of Triplasia has not been
studied by the present writers, and no quadrate topotypes have been
figured (those so placed by Beissel later being referred to a distinct
species). However, it is most probable that a large suite of speci-
mens would also show the quadrate character in this species. It is
probable that all quadrate species described in the past also have tri-
angular counterparts which in some cases have been described as
distinct species, although perhaps others have not yet been noted.

The present writers have examined very large suites of specimens
of the majority of species referred to both Triplasia and Frankeina
and have found that in most species there is a variation from straight
forms lacking the coil to specimens with a well-coiled base, and a
gradation from flattened uniserial portions (Flabellammina-like) to
triangular and quadrate tests. These gradations are more fully dis-
cussed under the specific descriptions, and are also illustrated on the
plates, but seem sufficient reason to suppress Centenarina, Tetraplasia,
and Frankeina all as synonyms of Triplasia.

ACKNOWLEDGMENTS

The writers have been aided immeasurably in this study in having
available many of the types of these species through the generosity of
Dr. C. I. Alexander, Magnolia Petroleum Corporation, San Antonio,
Tex., who presented his entire private collection of types to the U. S.
National Museum, including the many types of Frankeina described
by Alexander and Smith (1932). The large collection of Foraminifera
from Arkansas, made by the late Dr. W. H. Deaderick and bequeathed
to the National Museum, has supplied abundant specimens of many
old and new species.

Dr. Helmut Bartenstein, Deutsche Vacuum Oel, Celle, Germany,
generously supplied types of species described by Bartenstein and
Brand (1949 and 1951), some of which are here illustrated. Dr. R. W.
Harris, University of Oklahoma, lent the writers the types of the
Midway species from Arkansas, and Mrs. Billye Jobe, Humble Oil
and Refining Co., Tyler, Tex., presented to the National Museum
topotype specimens of T. fundibularis (Harris and Jobe).

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

To Cameron D. Ovey, British Museum (Natural History), we
are indebted for the fine suite of topotypes of the Recent species
Verneuilina variabilis Brady.

The remaining Lower Cretaceous specimens here described were
obtained by the writers in the course of preparation of a monographic
study of American Lower Cretaceous Foraminifera. These collections
were financed in part by a grant from the Penrose Bequest of the
Geological Society of America.

All illustrations were prepared by Mrs. Sally D. Lee, scientific
illustrator, Smithsonian Institution.

SYSTEMATIC DESCRIPTIONS
Family LITUOLIDAE
Genus TRIPLASIA Reuss, 1854

Genotype: Triplasia murchisoni Reuss, 1854.
Synonyms: Rhabdogonium Reuss, 1860 (part).
Verneuilina (part) Brady, 1884 (not d’Orbigny, 1840).
Haplophragmium (part) Andrae, 1890; Beissel, 1891; De Amicis,
1895; Liebus, 1911; Eichenberg, 1934; Keller, 1946 (not Reuss,
1860).
Ammobaculites (part) Franke, 1928 (not Cushman, 1910).
Frankeina Cushman and Alexander, 1929, and others.
Flabellammina (part) Alexander and Smith, 1932 (not Cushman,
1928).
Centenarina Majzon, 1948.
Tetraplasia Bartenstein and Brand, 1949.

Emended diagnosis—Test free, early portion may be planispiral,
later portion uniserial or may be uniserial throughout, uniserial por-
tion rhomboid in section, most commonly triangular, but occasionally
quadrate even in the same species ; sutures more or less arched on the
faces of the test, recurved at the angles; wall arenaceous ; aperture
terminal, occasionally on a neck, round to elongate. Range: Lower
Jurassic to Recent.

Remarks.—All species of Triplasia known to the writers are here
described, and figures are given for most of these. The earliest-known
species is from the upper Lias, and a few species are found in strata
of nearly every age from Lias to Recent. Apparently the genus had
two periods of climax, the Albian of America (seven species have
been described) and the Senonian (five species described from Amer-
ica and three others from Europe).

NO. 15

TRIPLASIA REUSS—LOEBLICH AND TAPPAN Uf

Geologic occurrence of Triplasia

Quaternary Recent T. variabilis
Pleistocene
Tertiary Pliocene T. wrighti
Miocene T. marwicki, T. minutwm
Oligocene T. hungarica, T. trigona
Eocene T. andraei
Paleocene T. fundibularis, T. sp.
Upper Cretaceous Danian
Maestrictian T. saratogensis
Senonian T. abkhasicus, T. beisseli, T. cushmani,
T. deadericki, T. plummerae, T. ro-
meri, T. rugosissima, T. taylorensis
Turonian T. murchisoni
Cenomanian T. nodosa
Lower Cretaceous Albian T. acutocarinata, T. glenrosensis, T.
goodlandensis, T. incerta, T. insolita,
T. rugosa, T. wenoensis
Aptian
Neocomian T. acuta, T. georgsdorfensis, T. gros-
serugosa, T. pseudoroemeri, T. mexi-
cana
Jurassic Malm T. commutata, T. elegans, T. jurassica
Dogger T. bartensteim
Lias T. kingakensis

TRIPLASIA KINGAKENSIS Loeblich and Tappan, new species
Plate 1, figures 1-8

Test large, elongate, early portion planispiral, with a compara-
tively large and well-developed coil of about 6 chambers, followed
by 4 or 5 uniserial and triradiate chambers, which increase gradually
in size in the megalospheric forms (figs. 3, 5-8) and flare more rapidly
in the microspheric form (figs. 1, 4a, b), occasional specimens de-
velop a somewhat weaker fourth angle, so that the test is irregularly
quadrate in section (figs. 2a, b), and in addition two specimens were
found which never developed the third angle, remaining in the Fla-
bellammina stage, whereas other specimens have a poorly developed
third angle, showing all gradations between; sutures moderately dis-
tinct, depressed, arched on the faces and curving downward at the
angles ; wall arenaceous, of medium to coarse particles, with consid-
erable cement, smoothly finished; aperture terminal, rounded, on a
slight neck.

Length of holotype, 1.53 mm.; greatest breadth, 0.94 mm. Length
of paratype of figure 6, 1.74 mm. ; greatest breadth, 0.73 mm. Length

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

of paratype of figure 3, 1.64 mm.; greatest breadth 0.52 mm. Other
specimens range from 0.91 to 1.69 mm. in length.

Number of specimens examined, 18.

Remarks.—This species resembles T. acutocarinata (Alexander and
Smith) in the flaring character of the test but differs in being smaller,
in having less-excavated sides and broader and more-rounded angles,
and in having a larger initial coil. It is quite variable in shape, rang-
ing from narrow to widely flaring and from flattened to triangular to
quadrangular in the later portion. The third and fourth angles are
less prominent than the two in the plane of the coil.

Occurrence.—Holotype (U.S.N.M. No. P266), figured paratypes
(U.S.N.M. Nos. P267a-g¢), and unfigured paratypes (U.S.N.M. No.
P268) all from the Kingak formation (Lower Toarcian) in a core at
2,028-2,048 feet in South Barrow Test Well No. 3, lat. 71° 09’ 4o” N.,
long. 156°34’45” W., south of Point Barrow, northern Alaska. This
species has a very restricted vertical range, as a complete core se-
quence throughout the Jurassic of this well showed the specimens to
be present only in this 20-foot interval.

TRIPLASIA BARTENSTEINI Loeblich and Tappan, new species
Plate 1, figure 9

Triplasia variabilis (Brady) BARTENSTEIN and Branp, 1937 (not Verneuilina
variabilis Brady, 1884), Abh. Senckenberg. naturf. Ges., No. 439, p. 185,
pl. 14A, fig. 6.

Description from Bartenstein and Brand (translation from Ger-
man) :

Test triangular in section, with rounded angles, greatest breadth in the later
portion. The early chambers form a small coil that becomes obscured by the
rapidly enlarging angles. Sutures but slightly marked, concealed by the coarsely
arenaceous surface of the test. Sides excavated. Aperture a slit at the thick-
ened point of intersection of the angles. Length 1.2 mm, breadth 0.9 mm.

Remarks.—This species differs from T. variabilis (Brady) in being
smaller and more flaring, with more deeply excavated sides and more
bluntly rounded angles. It is the only Middle Jurassic species of Tri-
plasia yet described.

Types and occurrence.—Holotype, specimen figured by Bartenstein
and Brand (Senckenberg Museum XX VII 644 a 2) from the Middle
Jurassic Dogger Epsilon (Wurttembergicus-Schichten) from the
well Bethel 2 at Bielefeld, Teutoberger Wald, in northwestern Ger-
many. This species was also recorded from the Parkinsoni-Schichten.

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 9

TRIPLASIA COMMUTATA (Loeblich and Tappan)
Plate 1, figures 12a-16
Frankeina sp. WICKENDEN, 1933, Trans. Roy. Soc. Canada, ser. 3, sect. 4, vol. 27,
Pp. iso, pit, ae, 2:

Frankeina commutata LorBLicH and Tappan, 1950, Journ. Washington Acad.
Sci., vol. 40, No. 1, p. 6, pl. 1, figs. 3a-4.

Test free, medium-sized, elongate, compressed and planispiral in
the early part, later uniserial and triangular in section, periphery
rounded; planispiral chambers strongly compressed, comprising a
relatively large portion of the test, about one and two-thirds volu-
tions in the coil; later portion uniserial, chambers increasing slowly
in diameter, in the holotype the final chamber being less in diameter
than the early coil, sides moderately excavated; sutures generally in-
distinct, in the coiled part straight or with a very slight backward
curve, very slightly depressed in the uniserial portion, but generally
indistinct except at peripheral angles; wall medium to coarsely are-
naceous, surface roughly finished ; aperture rounded, on a slight neck.

Length of holotype, 1.77 mm.; greatest diameter of coiled part,
0.75 mm.; greatest width of uniserial portion, 0.68 mm.; width from
center of side through opposite angle, 0.39 mm. Other specimens vary
from 0.65 mm. to 1.92 mm. in length, and in diameter of early coiled
part from 0.39 to 0.75 mm.

Remarks.—This species is closest to Triplasia incerta (Alexander
and Smith), from the Lower Cretaceous (Albian) of Texas, both
in size and in having a comparatively large coil. The Jurassic form
differs in having the pronounced coil of diameter equal to the breadth
of the later triangular portion, and in lacking the early Flabellammina
stage characteristic of T. incerta. The chambers do not enlarge as
rapidly, the sides are more excavated, and the sutures are more dis-
tinct in T. commutata.

Types and occurrence——Holotype (U.S.N.M. No. 106017), un-
figured paratypes (U.S.N.M. No. 106018), and unfigured hypotypes
(U.S.N.M. No. P878) from 94-99 feet above the base of the Rierdon
formation; unfigured paratypes (U.S.N.M. No. 106019) and un-
figured hypotypes (U.S.N.M. No. P879) from 99-104 feet above
the base of the Rierdon formation; unfigured paratypes (U.S.N.M.
No. 106020) from 134 feet above the base of the Rierdon formation ;
figured hypotype (U.S.N.M. No. P880) from 14-19 feet above the
base of the Rierdon formation (Callovian) ; all from the gorge of the
Shoshone River, 2.0 miles west of Cody, Park County, Wyo. Col-
lected by Ralph W. Imlay and Alfred R. Loeblich, Jr.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Figured paratype (U.S.N.M. No. 106021), unfigured paratypes
(U.S.N.M. No. 106022) and unfigured hypotypes (U.S.N.M. No.
P881) from 81 feet above the base of the Rierdon formation; un-
figured paratypes (U.S.N.M. No. 106023) and unfigured hypotypes
(U.S.N.M. No. P882) from 33 feet above the base of the Rierdon
formation ; unfigured hypotypes (U.S.N.M. No. P883) from 55 feet
above the base of the Rierdon formation; all from the southwest
corner of Red Dome, east of Bridger, in the Pryor Mountains, sec.
19, T. 7 S., R. 24 E., Carbon County, Mont. Collected by Ralph W.
Imlay and Alfred R. Loeblich, Jr.

Unfigured paratypes (U.S.N.M. No. 106024) and unfigured hy-
potypes (U.S.N.M. No. P884) from 47-52 feet above the base of the
Stockade Beaver shale (Callovian) ; figured hypotype (U.S.N.M. No.
P885) and unfigured paratype (U.S.N.M. No. P886) from 11-21 feet
above the base of the Redwater shale (Oxfordian) ; all from the
east side of Red Gulch, about 2.5 miles south of Little Big Horn River,
sec. 22, T. 58 N., R. 89 W., Sheridan County, Wyo. Collected by
Ralph W. Imlay and Alfred R. Loeblich, Jr.

Unfigured hypotypes (U.S.N.M. No. P887) from 7 feet above the
base of the Rierdon formation, in a gulch east of Swift Reservoir,
SW, sec. 6, T. 28 N., R. 10 W., Pondera County, Mont. Collected by
Ralph W. Imlay and Alfred R. Loeblich, Jr.

Unfigured hypotype (U.S.N.M. No. P888) from the Redwater
shale, 28-33 feet above the base, on the west side of Stockade Beaver
Creek, 5.0 miles northeast of Newcastle, sec. 18, T. 45 N., R. 60 W.,
Weston County, Wyo. Collected by Ralph W. Imlay and Alfred R.
Loeblich, Jr.

Unfigured hypotype (U.S.N.M. No. P889) from the Redwater
shale, 59-69 feet above base, 1.0 mile north-northeast of the center of
Spearfish, sec. 3, T. 6 N., R. 2 E., Lawrence County, S. Dak. Col-
lected by Ralph W. Imlay and Alfred R. Loeblich, Jr.

Unfigured hypotype (U.S.N.M. No. P8g90) from 1 foot above
the base; unfigured hypotypes (U.S.N.M. No. P891) from 9g feet
above the base; unfigured hypotype (U.S.N.M. No. P8&92) from 13
feet above the base; unfigured hypotypes (U.S.N.M. No. P8g3)
from 19 feet above the base; figured hypotype (U.S.N.M. No. P894)
from 22 feet above the base, and unfigured hypotypes (U.S.N.M. No.
P895) 25 feet above the base; all from the Swift formation (Oxford-
ian), 1.0 mile southwest of Landusky, sec. 32, T. 25 N., R. 24 E.,
Little Rocky Mountains, Phillips County, Mont. Collected by Ralph
W. Imlay and Alfred R. Loeblich, Jr.

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN Tey

TRIPLASIA ELEGANS (Mjatliuk)
Plate 1, figure 10

Frankeina elegans Mjatuiux, 1939, Neftianyi Geol. Razved. Inst., Trudy, ser.
A, vol. 120, pp. 48 (Russian), 71 (English), pl. 2, fig. 26.

Remarks.—This is a rather narrow species, with comparatively
small coil, closest in appearance to T. glenrosensis, new species, al-
though with a smaller coil, more excavated sides, and more depressed
sutures. The locality description referred to “isolated specimens,” but
the specific description refers only to the holotype.

Type and occurrence.—Hclotype (Geol. Oil Inst. collection, Lenin-
grad) from the Upper Jurassic, Lower Volga series, zone of Per-
isphinctes panderi d’Orbigny at a depth of 9-13 meters in Vostokneft
Well No. 1501, near the Station Ozinki on the Ryazan-Uralsk rail-
road, in the southern part of the Obschiy Syrt, Saratov District,
I S.5.h.

TRIPLASIA JURASSICA (Mjatliuk)
Plate 1, figures Ita, b

Flabellammina (Frankeina?) jurassica MJATLIUK, 1939, Neftianyi Geol. Razved.
Inst., Trudy, ser. A, vol. 120, pp. 47 (Russian), 70 (English), pl. 2, figs.
22 a-b.

Translation from Russian (p. 47), from Ellis and Messina (sup-
plement for 1950), states in part:

Test rounded-triangular, in the early stages close-coiled, later becoming
uniserial. . . . On one of the lateral surfaces of the test there is a slight ridge,
extending from the spire to the base of the last chamber. There is a correspond-
ing depression on the opposite side of the test... .

... the present species differs [from Flabellammina alexanderi Cushman]
in its less curved sutures and in the presence of a distinct costa, relating it to
the genus Frankeina. However, its cross-section not being triangular, we refer
it to the genus Flabellammina.

Length of holotype is given as 1.22 mm., width 0.66 mm., thickness
0.29 mm.

Remarks.—This species is very similar in character to the Lower
Cretaceous T. incerta, which also shows a close relationship between
Triplasia and Flabellammina. For this reason, and because the test
does develop a distinct third angle, we have here placed this species
in Triplasia.

Type and occurrence.— Holotype (Geol. Oil Inst. collection, Lenin-
grad) from the Upper Jurassic, Lower Volga series, zone of Per-
isphinctes panderi d’Orbigny, dark gray shaly clay interbedded with

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

dark brown and dark gray bituminous shales, with layers of dark
gray clay, at a depth of 22-24 meters (given on p. 48 as 22-29 meters),
in Vostokneft Well No. 1501, near the station Ozinki, on the Ryazan-
Uralsk railroad, southern part of the Obschiy Syrt, Saratov District,
U.S.S.R. Isolated specimens.

TRIPLASIA ACUTA Bartenstein and Brand
Plate 2, figure 6a, b

Triplasia emslandensis Bartenstein and Brand subsp. acuta BARTENSTEIN and
Branp, 1951, Abh. Senckenberg. naturf. Gesell., No. 485, p. 274, pl. 3,
fig. 68.

The original description states (translation from German):

Diagnosis: A subspecies of Triplasia emslandensis with sharp angles, unequal
strongly concave sides and almost completely suppressed coil.

Description: Test free, elongate, older chambers planispirally coiled, how-
ever the spire more or’less imperceptible, triangular with sharp angles and
variable strongly concave sides. Sutures somewhat depressed, strongly arched,
wall mostly medium-grained to finely arenaceous. Greatest breadth in the
younger part of the test due to slight and uniform enlarging. End chamber
truncate above, aperture irregularly elongate, situated on a small neck.

Length of holotype, 1.65 mm.

Remarks.—T. emslandensis subsp. emslandensis is a synonym for
T. georgsdorfensis (see discussion under that species). Therefore
the name T. emslandensis is invalid and if considered only a sub-
species, the present form would thus be known as T. georgsdorfensis
subsp. acuta. As the present species seems quite distinct, however, the
subspecific name acuta is here raised to specific rank.

Types and occurrence —Holotype (Senckenberg Natur Museums,
Frankfurt am Main collections) from Grenzbereich, Upper Valen-
dian 3 to 2, in Bohrung Diiste (Blatt 1736, neu 3317) K 521.6-526.3
m, northwestern Germany.

TRIPLASIA GEORGSDORFENSIS (Bartenstein and Brand)
Plate 1, figures 17a-21b

Tetraplasia georgsdorfensis BARTENSTEIN and Brawnp, 1949, Journ. Paleontol.,
vol. 23, No. 6, p. 672, text figs. 9a, b; 1951, Abh. Senckenberg. naturf.
Gesell., No. 485, p. 275, pl. 11, figs. 70-71.

Triplasia emslandensis emslandensis BaRTENSTEIN and Brawnp, 1951, _ ibid.,
p. 274, pl. 3, figs. 65-67.

not Haplophragmium murchisoni Reuss, BrEtsset, 1891, Abh. preuss. geol.
Landesanst., Berlin, n.s., No. 3, p. 15, pl. 4, figs. 2, 5, 9.

The following description is from Bartenstein and Brand (1949),
the original description of Tetraplasia georgsdorfensis :

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 13

Test free, elongate, regularly increasing in size. Earliest chambers rarely
recognizable but planispirally coiled (often the coiled part is only suggested by
the irregularity of the juvenile chambers), later chambers uncoiled, quadrangu-
lar, walls depressed with more or less rounded edges; sutures somewhat
depressed. Walls rather coarsely arenaceous. Aperture terminal, central, round
or somewhat elongate, ...

The original description of the triangular form, named Triplasia
emslandensis emslandensis by Bartenstein and Brand, states (trans-
lation from German) :

Diagnosis: A subspecies of the species Triplasia emslandensis n. sp. with
broad angles, sharply excavated sides, thick-set chambers and a uniform rate
of growth of isolated single chambers.

Description: Test free, elongate stout, older chambers spirally enrolled, gen-
erally very thick, younger chambers broader, triangular with concave sides and
+ broadly rounded angles. Test generally very coarse, straight or weakly
curved in growth, greatest breadth reached by the youngest chambers, aperture
circular or elongate in the center of the short tubular or produced end chamber.
Sutures depressed, walls agglutinated, medium- to fine-grained. Often mature
single chambers occur that are caused by especially strong constriction of the
sutures and ultimate separation of the chambers and whose membership in
this species may be assumed.

Two form groups may be distinguished: one form with thicker, although
distinct, spire and truncated end chamber (principal occurrence in upper
Valendian), another form with long acuminate end chamber lacking a spire
and tendency to separation of single chambers (principal occurrence in Lower
Hauterivian).

Length of type of Tetraplasia georgsdorfensis, 1.25 mm. ; length of
type (fig. 17) of Triplasia emslandensis emslandensis, 1.5 mm. Length
of small quadrate hypotype (fig. 18), 0.75 mm.; length of hypotype
of figure 20, 1.33 mm.; breadth, 0.83 mm.; length of hypotype (fig.
21), 1.61 mm., greatest breadth, 0.62 mm.

Number of quadrate specimens examined by the present writers,
two. We also have three triangular specimens originally referred to
Triplasia emslandensis emslandensis by Bartenstein and Brand.

Remarks.—T. georgsdorfensis is the genotype species for the genus
Tetraplasia Bartenstein and Brand. However, as stated by Bartenstein
and Brand, “close relationship to Triplasia is indicated by individuals
showing transitional characters.” As similar transitional forms are
found in nearly every species of Tviplasia the present writers be-
lieve the triangular and quadrangular forms to be congeneric and con-
specific as well, as gradations are so frequent in the various species.
Furthermore, T. georgsdorfensis and T. emslandensis emslandensis
are similar in other respects, as can be seen from the figures and
descriptions, and have almost identical geologic ranges, the triangular
forms being somewhat more abundant, as is true of all species yet ob-

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

served by the writers. The name Tetraplasia is thus suppressed as
a junior synonym of Triplasia, but as Tetraplasia georgsdorfensis was
the first specific name proposed it has priority over Triplasia emslan-
densis and the species thus becomes known as Triplasia georgsdorfensis
(Bartenstein and Brand).

Bartenstein and Brand included in their synonymy of Tetraplasia
georgsdorfensis the Upper Cretaceous Haplophragmium murchisom
Reuss of Beissel, 1891, as they noted that three quadrate specimens
were figured by Beissel with seven triangular forms. Comparison of
topotypes of Beissel’s form and topotypes of T. georgsdorfensis shows
these species to be distinct, and in fact the Upper Cretaceous form
has been described as Frankeina beisseli Marie (1941).

Types and occurrence-——Holotype of Tetraplasia georgsdorfensis
and types of Triplasia emslandensis emslandensis (Senckenburg Mu-
seum collections) all from the upper Valendian in a deep well, Georgs-
dorf 9, at 625 meters in depth, at Emsland, Germany. The quadrate
hypotype specimens here figured (U.S.N.M. Nos. P896a, b) are from
the upper Valendian in Georgsdorf 7 well, at a depth of 440 meters;
the triangular figured hypotype (U.S.N.M. No. P897) and unfigured
triangular hypotypes (U.S.N.M. No. P898) from the lower Hauteri-
vian (Noricus-Schichten) Zgl. Spiekerberg, Messtischblatt 1950,
Germany.

The triangular forms have been recorded from upper Valendian
to upper Hauterivian, from rare to common, and sporadically abund-
ant in the lower Noricus-Schichten. The quadrate forms are very
rare from upper Valendian to lower Hauterivian.

TRIPLASIA GROSSERUGOSA ten Dam
Plate 2, figures 5a-c

Triplasia grosserugosa TEN Dam, 1946, Journ. Paleontol., vol. 20, p. 571, pl. 87,
figs. 6a-c.

Original description :

Test free, elongate, triangular in section. Sides distinctly concave, angles
rounded. Initial chamber globular, fairly large, followed by 2 or 3 uniserial
chambers, slightly curved backward along the angles. Sutures slightly de-
pressed, faintly curved. Aperture terminal, circular. Wall very coarsely are-
naceous, rather roughly finished.

Dimensions.—Length, 1.65 mm.; width, 0.85 mm.; diameter of proloculum,
0.4 mm.

Remarks.—According to ten Dam, “This species is somewhat similar
to Triplasia roemeri Reuss, but differs in its concave sides and in its
smaller number of chambers.”

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 15

Types and occurrence.—Holotype in Netherlands Geological Sur-
vey, Haarlem, from the Hauterivian (Lower Cretaceous), near bound-
ary stone No. 849, in the Glanerheek near the village of Glanerbrug,
Overijsel Province, Netherlands.

TRIPLASIA MEXICANA Loeblich and Tappan, new species
Plate 2, figures 21-22b

Test free, flaring, triangular in section, sides excavated, angles
subacute; early chambers in a small planispiral coil which may be
somewhat obscure, later chambers uniserially arranged, low and broad,
up to nine in number, triangular in section; sutures obscured but
gently arched on the faces, recurved at the angles; wall moderately
coarsely arenaceous, with occasional large grains, rather smoothly
finished ; aperture terminal, rounded.

Length of holotype, 1.79 mm.; greatest breadth, 0.94 mm. Length
of figured paratype, 1.01 mm. ; greatest breadth, 0.55 mm. Other para-
types range in length from 0.57 to 2.08 mm.

Remarks.—The writers have examined 55 specimens of this species.
It is closest to the Albian species T. acutocarinata but is smaller, about
one-third to one-half as large, and the sides are less excavated. It is
sufficiently similar, however, to suggest that it may be the ancestral
form for the Albian species.

Types and occurrence.—Holotype (U.S.N.M. No. P1022), figured
paratype (U.S.N.M. No. P1023), and unfigured paratypes (U.S.N.M.
No. Pro24, and in the Paleontological Laboratory, Gerencia de Ex-
ploracién of Petroleos Mexicanos, Mexico City, and Instituto de
Geologia, Mexico City) all from 300 feet below the top of the Lower
Cretaceous (Hauterivian) Barril Viejo shale, Potrero Ovallos, in
the Sierra Hermanos, lat. 27°27’ N., long. 101°28’ W., Coahuila,
Mexico. Collected by R. W. Imlay.

TRIPLASIA PSEUDOROEMERI Bartenstein and Brand
Plate 2, figures 1ta-4b

Haplophragmium sp. (? n. sp.) EI1CHENBERG, 1934, Niedersachs. geol. Ver. Han-
nover, Jahrb. 26, p. 151, pl. 17, figs. 2a-b.

Triplasia pseudoroemeri BARTENSTEIN and Branp, 1951, Abh. Senckenberg.
naturf. Gessell., No. 485, p. 274, pl. 3, fig. 60, pl. 13, fig. 362.

Tetraplasia quadrata BARTENSTEIN and Branp, 1051, ibid., p. 275, pl. 3, fig. 72.

According to Bartenstein and Brand (translation from German) :

Test free, uniformly broad, without a coil, with a thickened proloculus, tri-
angular with convex to straight sides (and in the younger part of the test

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

occasionally also weakly concave) and bluntly rounded corners, few chambers,
broader than high (only the final chamber is higher than broad) and gently
curved, slightly depressed sutures. Wall finely agglutinated with less-coarse
components. Final chamber weakly acuminate to the elongate aperture, which is
somewhat eccentric in position.

Remarks: The older chambers are occasionally somewhat irregularly biserial
and indicate therefore the beginning of the spiral coiling. A true coil is appar-
ently not evident.

Eichenberg’s specimen (figs. 4a, b) is very similar to the type of
this species and was described as follows (translation from German) :
“Shell coarsely arenaceous, with interspersed dark green irregular
grains. Three chambers, the first two inflated, ovate; the third larger,
rounded in section, elongated, 3-sided. Aperture an elliptical opening.”

Bartenstein and Brand separated the very similar quadrate forms as
a distinct species (T. quadrata) (figs. 2a, b) referred to their genus
Tetraplasia. Their description and figures (which are here refigured)
seem to make it evident that the two “species” are the same, the tri-
angular specimens being more abundant, and the quadrate form very
rare. The description of the quadrate forms given by Bartenstein and
Brand follows (translation from German) :

Test free, uniformly broad, without a coil, in its place the older chambers
are somewhat irregularly biserial. Sides only very weakly concave, angles
broadly rounded, cross section therefore approximately rectangular. Chambers
broader than high, only the end chamber increasing in height, sutures slightly
depressed and gently curved, test broadly rounded at the base, the top weakly
acuminate with the indication of an apertural neck and more or less rounded

aperture. Walls agglutinated, of fine to medium coarseness. The cross section
of the test shows broader and narrower sides.

The holotype of T. pseudoroemeri (fig. 1) is 1.2 mm. in length,
the type of “Tetraplasia quadrata” Bartenstein and Brand (fig. 2) is
1.1 mm. in length, and the triangular juvenile specimen of Eichenberg
(fig. 4) is 0.97 mm. in length and 0.6 mm. in breadth.

Remarks.—The triangular and quadrate specimens referred to
above are similar in having an abortive coil of two large, inflated, ad-
jacent chambers at the base, followed by the angular later chambers
with nearly flat sides and broadly rounded angles, and a moderately
produced, rounded aperture. Both authors recorded their forms as
“very rare,” although T. pseudoroemeri was stated to be locally fre-
quent, and all are from the Neocomian of northern Germany. Eichen-
berg’s specimen was from the Hauterivian, and Bartenstein and Brand
recorded the quadrate form only from the upper Valendian, and the
triangular one from the upper Valendian to lower Hauterivian.

This species is not so coarsely arenaceous as the Netherlands

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN Wy

Hauterivian species, Triplasia grosserugosa ten Dam, and is slightly
larger, but it is otherwise similar in shape; abortive coil, nearly flat
sides and rounded angles, low early chambers and much elevated
pyriform final chamber. The two may be conspecific, but an examina-
tion of the types is necessary for confirmation of this.

Types and occurrence.—Holotype from the subsurface upper Val-
endian from Bohrung Georgsdorf 6 (Blatt 1 799, neu 3408), K 477
m, in northwestern Germany. Holotype of “Tetraplasia quadrata’”
from the subsurface upper Valendian in Bohrung Georgsdorf 7, at K
450 m, in northwestern Germany.

Triangular juvenile specitnen of Eichenberg, from the Tenuis zone
of the Hauterivian of Wenden on the Mittellandkanal, in northwestern
Germany.

TRIPLASIA GLENROSENSIS Loeblich and Tappan, new species
Plate 2, figures 7a-13b

Frankeina goodlandensis Cushman and Alexander, STEAD, 1951 (not Cushman
and Alexander, 1929), Texas Journ. Sci., vol. 3, No. 4, p. 580, pl. 1, fig. 6.
Test large, early portion planispiral, later uniserial and triangular
or quadrangular in section, 5 or 6 inflated planispiral chambers form-
ing a large coil, followed by 2 to 5 uniserial rectilinear chambers which
may be triangular in section with nearly flat sides and broadly rounded
angles (figs. 7a, b) or quadrangular (14 percent, as fig. 13), or may
lose the third angle so that the later portion is flattened and Flabellam-
mina-like (fig. 8) or the later chambers may lose the angularity and
become rounded (figs. 9, 11) ; sutures arched at the center of the faces,
lower at the angles, rather obscure, but sometimes slightly depressed ;
wall finely arenaceous, with considerable cement, very smoothly
finished ; aperture terminal, slitlike.

Length of holotype (fig. 7), 2.08 mm.; breadth, 0.83 mm. Length
of paratype of figure 8, 2.13 mm.; breadth, 0.78 mm. Length of quad-
rate paratype (fig. 13), 1.27 mm.; breadth, 0.44 mm. Other paratypes
vary in length from 0.83 to 2.24 mm. and in breadth from 0.42 to
1.04 mm.

Remarks.—This species resembles Triplasia commutata (Loeblich
and Tappan) in possessing a large coil and in having a nearly paral-
lel-sided later portion. It differs in having more-rounded angles, in
the smoother finish of the wall, and in the planispiral chambers being
inflated and subglobular instead of compressed and flattened. It is
very similar to T. georgsdorfensis (Bartenstein and Brand) but has
a much finer-textured wall, less-incised sutures and less-excavated
faces. Of the 190 specimens examined, 26 are quadrate in section,

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Occurrence.—Holotype (U.S.N.M. No. P8g9), figured paratypes
(U.S.N.M. Nos. Pgooa-f), and unfigured paratypes (U.S.N.M. No.
Poor) all from the Glen Rose formation, 520 feet below the top
in yellowish-gray Orbitolina-bearing marl, in a road cut on the east
side of U. S. Highway 281, 2.4 miles north of the junction of U. S.
Highway 281 and Texas Highway 46, Comal County, Tex. All
specimens collected by Alfred R. Loeblich, Jr.

TRIPLASIA GOODLANDENSIS (Cushman and Alexander)
Plate 2, figures 14a-20

Frankeina goodlandensis CUSHMAN and ALEXANDER, 1929, Contr. Cushman Lab.
Foram. Res., vol. 5, p. 62, pl. 10, figs. 1, 2—-ALEXANDER and SMITH, 1932,
Journ. Paleontol., vol. 6, No. 4, p. 307, pl. 47, fig. 8; Lozo, 1944, Amer. Midl.
Nat; voll 31, Now 3) ps 542) pl. 3) fig. 3:

Test small for the genus, early portion planispiral and compressed,
later rectilinear and triangular (figs. 15-20) or quadrangular (fig. 14)
in section (about 3 percent of the specimens), sides concave, the
test may flare rapidly (fig. 20) or have nearly parallel sides (figs. 15,
16), periphery subacute ; chambers numerous, about 5 in the coil, in-
creasing rapidly in size in the uniserial portion, although the first
few uniserial chambers may be of slightly less diameter than the coil,
chambers curve strongly backward at the angles, and the final cham-
ber may reach half the distance to the coil at the margins, sides
moderately excavated ; sutures somewhat obscure, but may be slightly
depressed in the later portion, straight and radiate in the coil, but
strongly recurved at the angles of the triangular portion, in many speci-
mens the position of the sutures suggested by the alignment of shell
fragments such as Inoceramus prisms, as they are nearly always placed
parallel to the sutures ; wall arenaceous, of medium-sized grains with
considerable cement, surface smoothly finished ; aperture rounded, at
the end of the somewhat produced final chamber.

Length of holotype, 1.0 mm.; breadth, 0.40 mm. Length of topo-
type of figure 19, 1.01 mm.; breadth, 0.42 mm. Length of quadrate
topotype (fig. 14), 0.68 mm.; breadth, 0.34 mm. Length of flaring
topotype of figure 20, 1.14 mm.; breadth of coil, 0.16 mm.; greatest
breadth of triangular portion, 0.65 mm. Length of topotype of figure
15, 0.52 mm. Length of topotype of figure 16, 0.62 mm. Length of
topotype of figure 18, 0.73 mm. Length of topotype of figure 17,
0.52 mm.

Remarks.—This was the genotype species of Frankeina, and of the
862 specimens examined, 25 (about 3 percent) were quadrate. It can

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN Ig

be distinguished from other species of the genus by its rather small
size, sharp angles, and very strongly arched sutures.

Occurrence.—Holotype (Cushman Coll. No. 12030) and paratypes
(Cushman Coll. Nos. 12031 and 12032) collected by C. I. Alexander,
and figured topotypes (U.S.N.M. Nos. Pgo2a-g) and unfigured topo-
types (U.S.N.M. Nos. Pg1o-Pg914) collected by Helen Tappan Loeb-
lich and Alfred R. Loeblich, Jr., all from the Goodland formation
(blue marls and chalky limestones) at the Lake Worth Dam, west of
Fort Worth, Tarrant County, Tex.

Plesiotype (U.S.N.M. No. Pots) figured by Alexander and Smith
and unfigured hypotypes (U.S.N.M. Nos. Pg16-P920) all from the
upper Goodland formation at “Cragin Knobs” on the old Stove Foun-
dry Road, 3.8 miles west of Montgomery Street, in Fort Worth, Tar-
rant County, Tex. Collected by Helen Tappan Loeblich and Alfred
R. Loeblich, Jr.

Unfigured hypotypes (U.S.N.M. No. Pg21) from the Goodland
formation in the east bank of North Fork of Mary’s Creek on the
Fort Worth—Weatherford highway, 11.5 miles west of Fort Worth,
Tarrant County, Tex. Collected by Helen Tappan Loeblich and
Alfred R. Loeblich, Jr.

Unfigured hypotypes (U.S.N.M. No. Pg22) from the Goodland
formation in the south bank of Clear Creek, 0.2 miles west of Farm
Road 425 bridge, 4.9 miles west of U. S. Highway 77 junction, Den-
ton County, Tex. Collected by Helen Tappan Loeblich and Alfred
R» Loeblich, Jr:

Unfigured hypotypes (U.S.N.M. No. Pg923) from the Goodland
formation in a high cliff on the east side of U. S. Highway 77, 3.4
miles north of the main road turning east to Marietta, SE} sec. 31,
T.6S., R. 2 E., Love County, Okla. Collected by Helen Tappan Loeb-
lich and Alfred R. Loeblich, Jr.

TRIPLASIA ACUTOCARINATA (Alexander and Smith)
Plate 4, figures I-9

Frankeina acutocarinata ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6,
No. 4, p. 307, pl. 47, figs. 1, 6—TappPaNn, 1943, Journ. Paleontol., vol. 17,
No. 5, p. 483, pl. 77, figs. 14a, b—Lozo, 1944, Amer. Midl. Nat., vol. 31,
No. 3, p. 542, pl. 1, fig. 7.

Test free, large, elongate, flaring, the early portion a planispiral
coil, later uniserial and triangular in section, occasionally quadrangular,
periphery angular in the early stages, later the angles are bluntly
rounded; chambers recurved at the angles, strongly arched on the

40) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

center of each face; sutures radial and indistinct in the coil, but may
be reflected by the slightly lobulate periphery, strongly arched on the
faces of the polygonal rectilinear portion and slightly depressed; wall
medium to coarsely arenaceous, but may be smoothly finished, fre-
quently a number of /noceramus prisms are incorporated into the
wall; aperture rounded to elongate, at the end of the slightly produced
final chamber.

Length of holotype, figure 6, 3.3 mm. ; breadth, 1.4 mm. Length of
topotype of figure 5, 2.67 mm.; breadth, 1.14 mm. Length of hypo-
type of figure 2, 2.94 mm.; breadth, 2.16 mm.

Remarks.—Of the 4,913 specimens examined, 30 specimens (0.6
percent) were found to be quadrate in section. This species is char-
acterized by its large size, extremely excavated sides, and narrow keel-
like angles.

Types and occurrence.—This extremely large species is quite abun-
dant in the Lower Cretaceous (Albian) of Texas and Oklahoma. It
ranges from the Kiamichi formation (Iredericksburg group) through
the Duck Creek, Fort Worth, Denton, and Weno formations ( Washita
group). Holotype, shown in figures 6a, b (U.S.N.M. No. P903), and
unfigured paratype (Cushman coll. 17842) collected by C. I. Alex-
ander ; topotype of figures 5a, b (U.S.N.M. No. Pgo04) and unfigured
topotypes (U.S.N.M. No. Pgo5)) collected by Helen Tappan Loeblich
and Alfred R. Loeblich, Jr.; all from the Duck Creek formation, in
the south bank of a small stream, about 15 feet north of the road,
0.1 mile east of the bridge, 0.9 mile east of Fink, Grayson County, Tex.

Hypotype (U.S.N.M. No. Pgo6) of figure 1 from the Kiamichi
formation, a 1.3-foot zone of yellow-gray clay marl, 4 feet below the
top of the formation, in a road cut 2.6 miles northeast of Mosheim
schoolhouse, on the northerly route from Mosheim to Valley Mills,
just west of the bridge over Hogg Creek, in the southern part of
Bosque County, Tex. Collected by Helen Tappan Loeblich and Alfred
Re Loeblichy Js.

Hypotype of figure 3 (U.S.N.M. No. Pgo07) from the basal 4.5
feet of the Duck Creek formation, alternating yellow marls and yellow-
white limestone beds, in a low, east-facing cliff of a small tributary
creek which flows north into Oliver Creek, just within the eastern
edge of Wise County, Tex. Collected by Helen Tappan Loeblich and
Alfred R. Loeblich, Jr.

Hypotype of figure 9 (U.S.N.M. No. Pgo8) from the basal 3 feet
of the exposure of the upper Denton formation, a dark blue and black
fissile clay in an exposure along the steep west bank of the north fork

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 21

of Noland’s River, about 100 feet south of the bridge on the Godley-
Joshua road, 1.4 miles northeast of Godley, Johnson County, Tex.
Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.
Hypotypes (U.S.N.M. Nos. goga-c) of figures 4, 7, and 8 from
the upper part of the Weno formation exposed in a road cut on the
Fort Worth-Mansfield road, 0.3 mile south of the bridge across
Sycamore Creek, southeast of Fort Worth, Tarrant County, Tex.
Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.
Hypotype of figure 2 (U.S.N.M. No. Pio25) from the Fort
Worth formation, in a low road cut near the bend of the road, in
the northeastern corner of sec. 28, T.8 S., R. 2 E., in Love County,
Okla. Collected by Helen Tappan Loeblich and Alfred R. Loeb-
leh, Jr.

TRIPLASIA INCERTA (Alexander and Smith)
Plate 3, figures 1-13

Frankeina incerta ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6, No. 4,
p. 308, pl. 47, fig. 4 -TApPPAN, 1943, Journ. Paleontol., vol. 17, No. 5, p. 483,
pl. 77, figs. 15, 16.

Test free, large, early portion planispirally coiled, flattened, later
portion uniserial and Flabellammina-like with arched flattened cham-
bers for a variable portion of the test (figs. 2, 11), later portion de-
velops a third angle, usually lower than the two in the plane of coiling,
and occasionally a fourth, so that in the later portion the test is
triangular (figs. 1-7, 9-13) or quadrangular (fig. 8) sides moderately
excavated, periphery subangular ; chambers numerous, about four in
the coil, later portion with about 8 uniserial chambers, increasing
very slowly in size so that sides are nearly parallel, chambers arched
at the center of the sides, bending downward at the angles; sutures
may be slightly depressed, but usually indistinct; wall coarsely are-
naceous, containing numerous shell fragments in a ground mass of
finer material, surface somewhat rough; aperture terminal, rounded
to elongate, at the end of a short blunt necklike extension of the
final chamber.

Length of holotype (fig. 7), 1.7 mm.; breadth, 0.95 mm. Length of
hypotype of figure 1, 4.08 mm. Length of hypotype of figure 2,
2.49 mm.; breadth, 0.77 mm.; thickness through Flabellammina
stage, 0.28 mm.; thickness through triangular portion, 0.34 mm.
Length of hypotype of figure 3, 2.78 mm. Length of hypotype of
figure 4, 3.07 mm.; length of Flabellammina stage, 0.83 mm. ; breadth,
0.68 mm. Length of hypotype of figure 5, 4.24 mm. Length of hypo-

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

type of figure 6, 2.55 mm. Length of quadrangular hypotype of figure
8, 1.61 mm.; breadth, 0.83 mm. Length of hypotype of figure 9,
2.70 mm. ; length of Flabellammina stage, 1.38 mm. ; breadth, 1.07 mm.
Length of hypotype of figure 10, 1.09 mm. Length of hypotype of
figure II, 1.14 mm.; breadth, 0.70 mm. Length of hypotype of figure
12, 1.51 mm. Length of hypotype of figure 13, 1.82 mm.

Number of specimens examined, 882, of which 30 (3.4 percent)
are quadrate.

Remarks.—This is an extremely variable species with a Flabellam-
muna portion of variable length (figs. 3, 11), the third angle develop-
ing late and somewhat lower than the other two angles. Occasional
specimens develop a fourth angle (fig. 8), and some with a triangular
test lose this third angle, and revert to a Flabellammina form (fig. 6).

Occurrence.—This species was first described from the Fort Worth
and Denton formations (Alexander and Smith, 1932) and later re-
corded from the Duck Creek formation (Tappan, 1943). The range
is here extended downward into the Kiamichi formation and upward
to the Weno formation.

Holotype, figure 7 (U.S.N.M. No. P924), and paratypes (Cushman
coll. No. 17850) collected by C. I. Alexander, from the upper Fort
Worth formation, in an exposure in the east bank of Denton Creek,
1.4 miles east of Justin, Denton County, Tex.

Hypotypes of figures 2 and 11 (U.S.N.M. Nos. Pg25a, b) from the
Duck Creek formation, from a 5.5-foot section of blue-gray shale,
about 30 feet above the base of the exposure, in the west bank of the
Red River, in SWi sec. 22, T. 8 S., R. 2 E., on the southwest side
of Horseshoe Bend, Love County, Okla. Collected by Helen Tappan
Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 8 (U.S.N.M. No. P926) from the Weno forma-
tion, in a road cut exposing about 10 feet of interbedded limestones
and yellow-gray marls, on the Fort Worth—Mansfield road, 0.3 mile
south of the bridge over Sycamore Creek, southeast of Fort Worth,
in Tarrant County, Tex. Collected by Helen Tappan Loeblich and
Alfred R. Loeblich, Jr.

Hypotype of figure 10 (U.S.N.M. No. P927) from a 3-foot section
of the dark gray and yellow shales of the Fort Worth formation,
about 10 feet above the base and 4 feet below the top of the exposure
in a deep road cut on the west side of U. S. Highway 75 (the Deni-
son-Durant road), 1,000 feet north of the Calvary Cemetery, 1.6 miles
north of the intersection with Main Street in Denison, Grayson

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 23

County, Tex. Collected by Helen Tappan Loeblich and Alfred R.
Loeblich, Jr.

Hypotypes of figures 6, 12, 13 (U.S.N.M. Nos. Pg28a-c) from the
upper Denton formation, about 7.5 feet of yellowish and reddish clay,
3 feet above the base and 5 feet below the top of the exposure, along
the steep west bank of the north fork of Nolands River, about 100
feet south of the bridge on the Godley-Joshua road, 1.4 miles north-
east of Godley, Johnson County, Tex. Collected by Helen Tappan
Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 9 (U.S.N.M. No. Pg29) from the Kiamichi
formation in a 3-foot exposure of bluish-gray clay and nodular lime-
stones, just above the Edwards limestone and immediately below the
Georgetown limestone (Duck Creek member) in the eastern bank of
Hogg Creek just north of the bridge at Patton, McLennan County,
Tex. Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 3 (U.S.N.M. No. P930) from the uppermost
2 feet of the Kiamichi formation, just below the Duck Creek forma-
tion in thin shaly layers between ledges of Gryphea shell agglomerate,
in a high, north-facing slope on the south bank of the Red River,
cleared by excavation at the site of the Denison Dam, north of Deni-
son, Grayson County, Tex. This slope is now covered and grassed
over. Collected in July 1940 by Helen Tappan Loeblich and Alfred
iz. Loeblich,jx.

Hypotype of figure 1 (U.S.N.M. No. P931) from the Fort Worth
formation, in a 6-foot exposure of yellowish-gray marls in a road
cut on the east side of U. S. Highway 77, 1.4 miles south of the south
end of the bridge across the Red River, in Cooke County, Tex. Col-
lected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 5 (U.S.N.M. No. P932) from the upper 4.5
feet of alternating yellow-gray limes and marly clays of the Fort
Worth formation, in a west-facing creek bank, just north of U. S.
Highway 70, and across the railroad tracks, in SW4NE} sec. 20, T.
6 S., R. 7 E., about 0.9 mile east of Aylesworth, Marshall County,
Okla. Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 4 (U.S.N.M. No. P933) from the Fort Worth
formation, from a 3-foot marl zone between heavy limestone beds,
10 feet above the base of a road cut in the 1700 block of East Lancaster
Street, just west of the corner of Riverside Drive and Lancaster
Street, in the Fort Worth—Dallas highway, in eastern Fort Worth,
Tarrant County, Tex. Collected by Helen Tappan Loeblich and Alfred
R. Loeblich, Jr.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

TRIPLASIA INSOLITA (Alexander and Smith)
Plate 5, figures 1a-7

Frankeina insolita ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6, No. 4,
p. 308, pl. 47, figs. 2, 3—TApPpPAN, 1943, Journ. Paleontol., vol. 17, No. 5,
p. 483, pl. 77, figs. 17a-18b.

Test free, large, early portion planispiral and compressed, then
becoming Flabellammina-like for a short distance (figs. 2a, b), or
abruptly triangular (figs. 1a, b), or quadrangular (figs. 3a, b), sides of
test somewhat excavated, angles bluntly rounded, chambers numerous,
at first increasing rapidly in size from the four or five of the tiny
planispiral coil, later chambers increase more slowly in size, so that
sides of test are nearly parallel; sutures obscure, somewhat more
distinct in the planispiral portion; wall coarsely arenaceous, incorpo-
rating many shell fragments, such as Inoceramus prisms; aperture
terminal, rounded.

Length of holotype (fig. 1), 2.7 mm.; breadth, 0.9 mm. Length
of topotype of figure 2, 2.60 mm.; breadth, 1.07 mm.; thickness in
early portion of test, 0.28 mm.; thickness near aperture, 0.62 mm.
Length of topotype of figure 3, 2.00 mm.; breadth, 1.20 mm.; thick-
ness, 1.01 mm.

Number of specimens examined, 328.

Remarks.—This is a large and extremely variable species and
shows all gradations between the flattened forms, the triangular and
the quadrangular ones.

Types and occurrence.—Although Alexander and Smith state
(1932, p. 308) that “F. insolita has been found in samples from the
Duck Creek formation only, and is thus an excellent marker of basal
Washita,” the writers have found specimens in the Fort Worth,
Weno, and Main Street formations which appear identical in all re-
spects to typical T. insolita. It seems probable that this species is
characteristic of a limy lithologic facies, and is found where these
formations are of similar facies as the Duck Creek limes and marls.

Holotype, figure 1 (U.S.N.M. No. P934), and paratypes (U.S.N.M.
No. P935 and Cushman Coll. 17849) collected by C. I. Alexander ;
and topotypes of figures 2 and 3 (U.S.N.M. Nos. Pg36a-b) and un-
figured topotypes (U.S.N.M. No. P937) collected by Helen Tappan
Loeblich and Alfred R. Loeblich, Jr.; all from the Duck Creek forma-
tion, in a low north-facing cliff on the south bank of a small stream
north of the road, 0.1 mile east of the bridge, 0.9 mile east of Fink,
Grayson County, Tex.

Hypotype of figures 5 and 7 (U.S.N.M. Nos. Pg38a, b) from the

NO. 15 TRIPLASIA REUSS—-LOEBLICH AND TAPPAN 25

Main Street formation, in 5.5 feet of section of alternating limes and
marls (Exogyra arietina zone), 4 feet below the top of the section
exposed in a road cut on the road leading eastward to Grayson Bluff,
about 1 mile east of the Fort Worth—Denton highway, 3.5 miles north-
east of Roanoke, Denton County, Tex. Collected by Helen Tappan
Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 6 (U.S.N.M. No. P939) from the Fort Worth
formation, in the lowest 5.5 feet of alternating limestones and light
gray marls exposed in a road cut on the east side of U. S. Highway
77, 1 mile west-southwest of the Gainesville courthouse square, just
south of a small bridge, in Cooke County, Tex. Collected by Helen
Tappan Loeblich and Alfred R. Loeblich, Jr.

Hypotype of figure 4 (U.S.N.M. No. Po40) from the alternating
thin limestones and thicker marl beds of the upper Weno formation,
in the upper one-half of the lower 5.5 feet exposed, and just under
a 1-foot limestone bed, 7 feet below the top of the formation, in a road
cut on the west side of the Fort Worth—Burleson highway, where it
swings southward near the top of the hill, 0.25 mile southeast of the
fork of the Fort Worth-Everman road from the Fort Worth-Burleson
highway, southeast of Fort Worth, Tarrant County, Tex. Collected
by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.

TRIPLASIA NODOSA Loeblich and Tappan, new species
Plate 5, figures 8-15

Test free, narrow, elongate, base rounded and bulbous, chambers
uniserially arranged, varying from somewhat flattened to distinctly
triangular or quadrangular in section, or even biformed, flattened at
first (fig. 11) and developing the third angle late in its growth, or
triangular at first and losing the third angle in the later development
(fig. 10), the later chambers sometimes inflated and nodular in ap-
pearance and separated by definite constrictions (figs. 13, 14) ; sutures
obscure to strongly constricted; wall coarsely arenaceous, incorpo-
rating grains and shell fragments of varying sizes, surface very rough
and irregular ; aperture rounded at the end of a narrow, much-pro-
duced neck.

Length of holotype (fig. 12), 1.27 mm.; breadth, 0.60 mm. Length
of paratype of figure 8, 0.96 mm.; breadth, 0.49 mm. Length of
paratype of figure 10, 1.53 mm.; breadth, 0.70 mm. Length of para-
type of figure 11, 1.51 mm.; breadth, 0.60 mm. Length of paratype
of figure 13, 1.22 mm.; breadth, 0.44 mm.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Remarks.—This extremely variable species is closest in appearance
to the early Washita Triplasia rugosa (Alexander and Smith), but
differs in the much more elongate neck, absence of a distinct coil, more
nodular appearance of the chambers and greater constriction of the
sutures. The later chambers are generally widest at their base and
taper sharply toward the aperture.

Types and occurrence.—Holotype (fig. 12, U.S.N.M. No. Pg4r),
paratypes of figure 9 and 10 (U.S.N.M. Nos. Po42a-b), and unfigured
paratypes (U.S.N.M. No. P943) all from the Grayson formation,
basal 3 feet, just above the Main Street formation, in a road cut on the
south bank of Chuckwa Creek, 1.0 mile north of Durant, on U. S.
Highway 75, in NE} sec. 29, T.6 S., R. 9 E., Bryan County, Okla.
Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr., 1937.

Paratypes of figures 8, 14, 15 (U.S.N.M. Nos. Po44a-c) and un-
figured paratypes (U.S.N.M. No. P945) all from the Grayson forma-
tion, at Grayson Bluff, a high southwest-facing bluff on Denton Creek,
3.5 miles northeast of Roanoke, Denton County, Tex. Collected by
Helen Tappan Loeblich and Alfred R. Loeblich, Jr., 1937.

Unfigured paratypes (U.S.N.M. No. P946) from the Del Rio clay,
15.5 to 21 feet above the base exposed, on the west bank of Shoal
Creek, just south of the bridge at 34th Street and just north of a
fault, in Austin, Travis County, Tex. Collected by Helen Tappan
Loeblich and Alfred R. Loeblich, Jr., July 1940.

Paratypes of figure 13 (U.S.N.M. No. P947) and unfigured para-
types (U.S.N.M. No. P948) from thin marl beds between limestone
ledges of the Main Street formation, basal 5.5 feet of section exposed
at an underpass of the Atchison, Topeka and Santa Fe Railroad, on
the Cleburne-Hillsboro road, just south of the city of Cleburne, in
Johnson County, Tex. Collected by Helen Tappan Loeblich and Alfred
R. Loeblich, Jr., July 1940.

Paratype of figure 11 (U.S.N.M. No. P949) and unfigured para-
type (U.S.N.M. No. Pg50) from the thin marl seams between project-
ing limestone ledges of the Main Street formation, in an 8-foot, west-
facing bank of a small stream, east of the road leading south one
block east of the eastern edge of the campus (Fort Worth—Crowley
road), 3.9 miles south of the southeastern edge of the Baptist Semi-
nary Campus, south of Fort Worth, Tarrant County, Tex. Collected
by Helen Tappan Loeblich and Alfred R. Loeblich, Jr., July 1940.

Unfigured paratypes (U.S.N.M. No. P951) from the lower Main
Street formation, in thin marl seams between heavy beds of lime-
stone, in a road cut on the north side of the road, at the western edge

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 27

of the Federal Narcotic Farm, southeast of Fort Worth, Tarrant
County, Tex. Collected by Helen Tappan Loeblich and Alfred R.
Loeblich, Jr., June 1939.

TRIPLASIA RUGOSA (Alexander and Smith)
Plate 3, figures 14-20c

Flabellammina rugosa ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6,
No. 4, p. 302, pl. 45, figs. 6-7 (not 8-9), text figs. 1, 2c (not 2b).—Tappan,
1943, Journ. Paleontol., vol. 17, No. 5, p. 482, pl. 77, figs. Ila-13.

not Flabellammina rugosa Alexander and Smith, CusuMaAn, 1946, U. S. Geol.
Sury. Prof. Pap. 206, p. 24, pl. 4, figs. 9-10.

Test free, of medium size, elongate, either with a bulbous base
(fig. 18) or an obscure coil (figs. 19, 20), later compressed (fig. 20) to
subtriangular (fig. 19) or quadrangular (fig. 16) in section; chambers
of the early coil usually somewhat obscure (fig. 19), those of the
early uncoiled portion slightly arched, later chambers irregular in
appearance, of somewhat greater breadth than height, final chamber
may be somewhat pyriform in outline (figs. 16, 18, 19) ; sutures gen-
erally indistinct, slightly arched, later ones somewhat constricted and
nearly straight; wall coarsely arenaceous, incorporating various-sized
grains and Inoceramus prisms, surface roughly finished; aperture at
the end of a short neck, rounded to somewhat elongate.

Length of holotype, 1.61 mm.; breadth, 0.78 mm.; thickness,
0.34 mm. Length of hypotype of figure 16, 1.82 mm.; breadth,
0.55 mm. Length of hypotype of figure 15, 1.46 mm.; breadth,
0.57 mm. Length of topotype of figure 17, 1.38 mm.; breadth, 0.64
mm. ; thickness, 0.37 mm. Length of topotype of figure 18, 1.53 mm. ;
breadth, 0.62 mm. Length of topotype of figure 19, 1.16 mm. ; breadth,
0.38 mm.

Remarks.—tThis species was originally described as a species of
Flabellammina and the holotype is somewhat flattened. Nevertheless,
because the majority of specimens are much less noticeably flattened,
but are triangular to quadrate, the species is here considered to be-
long to Triplasia. It is similar in some respects to T. incerta in that
there may be an elongate flattened portion before the test becomes
triangular or quadrangular.

Alexander and Smith, and later Cushman, also referred Upper
Cretaceous specimens to this species. However, their specimens from
the Austin chalk are typical Flabellammina and may possibly belong
to Flabellammina clava Alexander and Smith, which was also de-
scribed from the Austin chalk, and is of similar size, shape, and ap-

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

pearance. The Austin chalk specimens are much more compressed
than the present species, with broader and lower chambers and with
strongly arched sutures. They also lack the apertural neck of T.
rugosa.

Types and occurrence.—Holotype (fig. 20, U.S.N.M. No. Pg52)
and unfigured paratype (U.S.N.M. No. P953) collected by C. I.
Alexander, from the lower Duck Creek limestone and marl; and
topotypes of figures 17-19 (U.S.N.M. No. P954a-c) collected by Helen
Tappan Loeblich and Alfred R. Loeblich, Jr., June 1939, from the
basal 1-foot of alternating limestones and marls of the Duck Creek
formation (Lower Cretaceous, Albian), all from the south bank of
a small creek, about 15 feet north of the road leading eastward from
Fink, 1 mile east of town, in Grayson County, Tex.

Hypotype of figure 14 (U.S.N.M. No. Pg55) from the blue-gray
shales of the Duck Creek formation, from 19 to 24 feet above the
base, and hypotypes of figures 15 and 16 (U.S.N.M. Nos. Pg56a-b)
from 24 to 30 feet above the base, on the west bank of the Red River,
in SWi sec. 22, T.8S., R. 2 E., on the southwest side of Horseshoe
Bend, Love County, Okla. Collected by Helen Tappan Loeblich and
Alfred R. Loeblich, Jr., August 1939.

We have recorded this species from many Duck Creek outcrops in
Texas and Oklahoma, as well as from the lower Fort Worth lime-
stone of Oklahoma.

TRIPLASIA WENOENSIS (Alexander and Smith)
Plate 4, figures 10-15b

Frankeina wenoensis ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6,
No. 4, p. 300, pl. 47, fig. 5.

Test free, small for the genus, flaring, early portion planispiral,
later uniserial and triangular or very rarely quadrate (fig. 15), sides
moderately excavated, angles bluntly rounded; chambers few in num-
ber with only 3 or 4 comprising the uniserial portion, rather broad and
low ; sutures indistinct, occasionally marked by slight indentations at
the angles of the test, arched across the center of each side; wall
arenaceous, of medium-sized grains in a finer ground mass, very
roughly finished ; aperture terminal, rounded, on a short blunt neck.

Length of holotype (fig. 14), 1.30 mm.; breadth, 0.78 mm. Length
of hypotype of figure 10, 0.91 mm.; breadth, 0.57 mm. Length of
hypotype of figure 12, 0.81 mm.; breadth, 0.62 mm. Length of hypo-
type of figure 13, 1.51 mm.; breadth, 0.99 mm. Length of quadrate

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 29

hypotype of figure 15, 1.64 mm.; breadth, 0.78 mm. Other hypo-
types range from 0.83 to I.14 mm. in length.

Remarks.—Alexander and Smith stated that “This species is re-
stricted to the Weno formation, but the fact that it is a rather rare
form detracts from its value as an horizon marker. The writers’ col-
lections have failed to disclose a locality where individuals occur in
abundance.” Actually, the species was based on a single specimen, the
holotype, which was the only specimen in the Alexander collection
(now in the U.S. National Museum). In the large number of Washita
samples examined by the present writers, this species is very rare,
being represented by a total of eight hypotype specimens. However, it
has been found in the Main Street and Georgetown formations, as
well as the Weno formation, and its range within the middle Washita
was apparently governed by the environment, as all specimens occur
in the marly limestone facies of the respective formations. In spite
of its rarity, this species is represented by two quadrate specimens,
which are otherwise similar to the more typical triangular forms.

This species resembles T. acutocarinata (Alexander and Smith) in
the somewhat flaring character but is smaller and has less-excavated
sides, more-rounded angles, and more-obscure sutures.

Types and occurrence.—Holotype (U.S.N.M. No. P957) from the
Weno formation (Lower Cretaceous, Albian) in a roadside ditch on
the Fort Worth-Mansfield road, 0.25 mile south of the bridge over
Sycamore Creek, Tarrant County, Tex. C. I. Alexander collection.

Hypotype of figure 13 (U.S.N.M. No. P958) from the upper 10
feet exposed of the interbedded light yellow-gray marls and lime-
stones of the Weno formation in a road cut on U. S. Highway 287
(Fort Worth-Mansfield road), 0.3 mile south of the bridge over
Sycamore Creek, southeast of Fort Worth, in Tarrant County, Tex.
Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.,
August 1941.

Hypotypes of figures 10-12 and 15 (U.S.N.M. Nos. Pg5ga-d),
from the lower 1-foot marl bed of the Weno formation, exposed
beneath a heavy limestone and about 15-20 feet stratigraphically be-
low the Paw Paw formation, in a road cut on the east side of the
old Mansfield road, downhill and to the north of the Paw Paw ex-
posure, 0.25 mile south of the bridge over Sycamore Creek, west of
the Glen Garden Country Club, 3 miles southeast of Fort Worth,
Tarrant County, Tex. Collected by Helen Tappan Loeblich and
Alfred R. Loeblich, Jr., October 1943.

Because of the many changes in the highways over the past 20

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

years, it is uncertain which of the two preceding localities of the
present writers represents the actual type locality of Alexander and
Smith. They are only a fraction of a mile distant, however.

Unfigured hypotype (U.S.N.M. No. P960) from the upper 3 feet
exposed of the alternating limestones and light gray marls and yel-
lowish and brownish clays of the lower Weno formation, about 8
feet above the contact with the Denton formation, on the south bank
of a tributary to Sycamore Creek, which was dammed to form Katy
Lake, 0.25 mile east of and below the dam, southeast of Fort Worth,
Tarrant County, Tex. Collected by Helen Tappan Loeblich and Alfred
R. Loeblich, Jr., September 1938.

Unfigured hypotype (U.S.N.M. No. Po961) from the upper 5 feet
exposed of the Main Street formation (Lower Cretaceous, Albian)
consisting of marl beds between limestone ledges, at an underpass
under the Atchison, Topeka and Santa Fe railroad just south of
Cleburne, on the Cleburne-Hillsboro road, in Johnson County, Tex.
Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.,
July 1940.

Unfigured hypotype (U.S.N.M. No. P962) from the Georgetown
formation (Lower Cretaceous, Albian), in marls between large fucoid-
bearing limestone ledges, containing Macraster elegans (Shumard),
along Smith Branch, the first main creek east of the town of George-
town, approximately one-half mile northeast (downstream) from
Texas State Highway 104, Williamson County, Tex. Collected by
Helen Tappan Loeblich and Alfred R. Loeblich, Jr., 1940.

TRIPLASIA MURCHISONI Reuss
Plate 5, figures 16-17¢c

Triplasia murchisom Reuss, 1854, Denkschr. Akad. Wiss. Wien, math.-naturw.
K1., vol. 7, p. 65, pl. 25, figs. 1a-2.

Ammobaculites variabilis (Brady) FRANKE, 1928 (not Brady, 1884), Abh.
preuss. geol. Landesanst., Berlin, n.s., No. 111, p. 166, pl. 15, figs. 6a-b.
not Triplasia murchisoni Reuss, Eccrr, 1899, Abh. bayer. Akad. Wiss., Miinchen,

K1. 2, vol. 21, pt. 1, p. 42, pl. 15, fig. 24.

Original description (translation from German) :

Up to 3.2 mm. high, inverted ovate, more or less elongate, base bluntly pointed,
quite sharply triangular, the lateral sides indented along the direction of the
long axis, so that the cross section presents an equilateral, quite sharply angled
triangle, with concave sides.

Chambers up to 10, increasing gradually in size from base to top, triradiate,
attached across their entire breadth with no separating constrictions; each
somewhat overlapping the preceding. The sutures only slightly depressed and

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 31

descending somewhat obliquely from the middle of each side to the angles of
the test. The last chamber elongated in a short, somewhat thickened, central
tubular neck, which carries on the apex the round bare aperture. The outer
surface of the shell rough.

Remarks.—All attempts by the writers to obtain specimens of this
species have been futile. However, Cushman stated (1948, p. 106)
that a study of type material in Vienna showed that the microspheric
generation possessed a coil, which was not present in the megalospheric
form.

This specimen figured by Egger is quite distinct, being much longer
and narrower, with higher chambers and horizontal and constricted
sutures. It does not even appear to belong to Triplasia, but only an
examination of the types could prove this.

Franke referred to Ammobaculites variabilis (Brady) some rapidly
flaring specimens with a triangular section and low, broad chambers.
They seem closer in appearance to T. murchisoni Reuss which is also
from the Senonian (or Turonian) in this general region. Typical
T. variabilis is larger, has a less flaring test which becomes nearly
parallel-sided in the later portion, and has a more slitlike aperture.

Occurrence.—Described from the Turonian, or lower Senonian,
Gosaugebilde, Mergel, present at Edelbachgraben, and very rare
at Wegschiedgraben in Gosau, Ostalpen (Salzburg), Austria.

TRIPLASIA ABKHASICUS (Keller)
Plate 7, figure 15

Haplophragmium abkhasicus KELier, 1946, Bull. Soc. Natur. Moscow, N.s.,
vol. 51 (Sect. Géol., vol. 21), No. 3, pp. 89 (Russian), 106 (English), pl. 1,
fig. 17; pl. 3, fiz, 10:

_ Translation from the Russian, p. 89 (Ellis and Messina, supple-
ment for 1950):

Test very large, tightly coiled in the initial portion, later becoming uniserial.
Width of the test in the initial portion about the same as in the final portion.
In transverse section the uniserial portion of the test is angular, almost tri-
angular, with a rounded dorsal and a somewhat tapering ventral side. In the
spiral portion of the test there are 4 or 5 chambers, forming a single volution.
Chambers of the uniserial portion number 2-4, sutures linear, slightly depressed.
Aperture poorly defined, apparently complex, terminal. Wall agglutinated, con-
taining prisms of Inoceramus, Pythonellas and calcareous fragments.

The type specimens measured 2.0 mm. in length, 0.72 mm. in width,
and 1.32 mm. in length and 0.70 mm. in width.
Remarks—According to Keller, the species is related to Haplo-

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

phragmium aequale Roemer, but differs in having slightly depressed
sutures and narrower uniserial chambers. It also seems close to
Rhabdogonium rémeri Reuss, from the German upper Senonian, but
has a better-developed coil. Possibly Reuss’s type is megalospheric
and Keller’s microspheric. Both species are “very rare.”

Types and occurrence—Types in the State Institute of Sciences,
Moscow, from the lower Senonian (Campanian), Upper Cretaceous,
on the southern flank of the Dzykhrin anticline on the River Mzymta,
environs of Sotchi, western Caucasus, Krasnodar, U.S.S.R.

TRIPLASIA BEISSELI (Marie)
Plate 5, figures 18-20b; text figures 1-10

Haplophragmium murchisoni (Reuss) BeEtssEL, 1891 (not Reuss, 1854), Abh.
preuss. geol. Landesanst., Berlin, n.s., No. 3, p. 15, pl. 4, figs. I-1o.

Ammobaculites murchisoni Beissel (not Reuss) FRANKE, 1928, Abh. preuss.
geol. Landesanst., Berlin, n.s., No. 111, p. 165, pl. 15, fig. 5.

Frankeina beisseli Martz, 1941, Mém. Mus. Nat. Hist. Natur., Paris, n.s., vol. 12,
No. I, p. 23, pl. 2, fig. 12a-c.

not Frankeina cushmani ALEXANDER and SMITH, 1932, Journ. Paleontol. vol. 6,
No. 4, p. 300, pl. 47, figs. 10, II.

not Tetraplasia georgsdorfensis BARTENSTEIN and Branp, 1949, Journ. Paleontol.
vol. 23, No. 6, p. 672, text figs. ga, b.

Test free, large, flaring, with a well-developed planispiral coil,
followed by a few triangular or quadrate uniserial chambers, sides
moderately excavated, angles broadly rounded; sutures distinct,
slightly depressed, gently arched on the faces, recurved at the angles ;
wall coarsely arenaceous, but grains of nearly uniform size, little
cement, so that the surface is rough in appearance ; aperture a terminal
slit, sometimes slightly produced.

Length of hypotype (fig. 18), 2.70 mm. ; breadth, 1.30 mm. Length
of hypotype of figure 19, 3.17 mm.; breadth, 1.48 mm. Length of
hypotype of figure 20, 1.61 mm.; breadth, 0.62 mm. Other hypotypes
range from 1.30 to 2.68 mm. in length.

Remarks—Although referred to Haplophragmium murchisoni
(Reuss) by Beissel, it differs from Reuss’s species in being less
flaring, in having a more-prominent coil and less-acute angles, in
lacking the neck and in having a slitlike rather than a rounded
aperture.

Beissel’s reference was also placed in synonymy of Frankeina
cushmani Alexander and Smith by the authors of that species, but the
European species is larger and more flaring and has better-defined and

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 33

slightly excavated sutures and a less coarsely arenaceous and more
smoothly finished wall.

Bartenstein and Brand (1949, p. 672) considered the quadrate
specimens figured by Beissel to belong to their new genus and species
Tetraplasia georgsdorfensis. The latter does not have as prominent
a coil, has less highly arched sutures and more excavated sides, and
is a much smaller species.

Types and occurrence—Holotype in the Marie collection, from
the Campanian (Upper Cretaceous), Belemnitella mucronata chalk
at Montereau, Seine-et-Marne, Paris Basin, France. Paratypes

Fics. 1-10.—Triplasia beisseli (Marie): 1, A biformed individual, which is
flattened in part and develops a third angle in the central portion of the test. 2,
Face view of a quadrate specimen. 3, Side view of a triangular specimen. 4,
Face view showing arched sutures. 5, Apertural view of a quadrate specimen.
6. Apertural view of a triangular specimen. 7, Opening of the penultimate cham-
ber. 8, Opening of one of the earliest uncoiled chambers. 9, A more excavated
quadrate specimen. 10, Sectional view. (After Beissel.)

(d’Orbigny collections, Muséum Nationale d’Histoire Naturelle,
Paris) from Meudon, Seine-et-Oise, France.

Hypotype of figure 18 (U.S.N.M. No. P963), hypotype of figure
19 (U.S.N.M. No. P964), and unfigured hypotype (U.S.N.M. No.
P965) all from the Mucronaten-Kreide, Friedrichsberg bei Aachen,
Germany (topotypes of Haplophragmium murchisoni of Beissel,
and identified as such by A. Franke), in the Alexander collection
deposited in the U. S. National Museum.

Hypotypes (U.S.N.M. No. P966) from the lower Senonian, of
Westphalia, Germany. Collected by A. Franke and donated to the
U. S. National Museum by C. I. Alexander.

Quadrate hypotype of figure 20 (U.S.N.M. No. P967) and un-
figured hypotypes (U.S.N.M. No. P968) from the lower Senonian,
between Hannover and Hildesheim, Germany. Collected by A. Franke
and donated to the U. S. National Museum by C. I. Alexander.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

TRIPLASIA CUSHMANI (Alexander and Smith)
Plate 7, figures Ia-5

Frankeina cushmani ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6,
No. 4, p. 300, pl. 47, figs. 10-11—CusHMAN, 1946, U. S. Geol. Surv. Prof.
Pap. 206, p. 25, pl. 4, figs. II, 12.

Test free, of medium size for the genus, flaring in the early portion,
later with nearly parallel margins, planispiral coil large, later uni-
serial portion triangular to quadrate with moderately excavated sides
and acute angles ; chambers low and broad, moderately arched on the
sides; sutures obscure, discernible largely by the alignment of the
larger grains and calcite prisms in the wall, rarely slightly depressed ;
wall coarsely arenaceous, with large grains and shell fragments in a
finer ground mass, roughly finished; aperture a terminal slit, not
produced.

Length of holotype (fig. 2), 1.51 mm.; breadth, 0.86 mm. Length
of hypotype of figure I, 1.43 mm.; breadth, 0.81 mm.; thickness,
0.52 mm. Length of quadrate hypotype of figure 3, 2.24 mm. Length
of hypotype of figure 4, 0.86 mm. Length of hypotype of figure 5,
0.99 mm.

Remarks.—Of the 232 specimens examined, 4 are quadrate, and
the remainder are triangular in section, a percentage of 1.72 quadrate
forms.

This species is characterized by the comparatively large coil, paral-
lel sides, and obscure sutures, and the coarsely arenaceous and roughly
textured wall. Alexander and Smith placed in the synonymy of this
species Haplophragmium murchisom (Reuss) of Beissel, 1891 (not
Triplasia murchisoni Reuss, 1854). Topotypes of Beissel’s form, |
identified by A. Franke, show that this is a distinct species (described
elsewhere in this paper as T. beisseli (Marie) ). T. beisseli is a larger
and more flaring species, with better-defined sutures and less coarsely
arenaceous and more smoothly finished wall.

Types and occurrence.—Holotype (fig. 2, U.S.N.M. No. Po89),
paratype figured by Alexander and Smith (U.S.N.M. No. Pg90),
and unfigured paratype (Cushman coll. No. 17852) all from the
Pecan Gap (Upper Cretaceous), in an abandoned clay pit, 1.0 mile
east of Farmersville, on the Farmersville-Greenville road, Collin
County, Tex. Collected by C. I. Alexander.

Hypotype of figure 3 (U.S.N.M. No. Pog1) and unfigured hypo-
types (U.S.N.M. No. Pg92) from the Upper Cretaceous Annona
chalk, about 1 mile north of the quarry of the Arkansas Lime Prod-
ucts Co., on Little River, Ark. Collected by W. H. Deaderick.

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 35

Hypotype of figure 1 (U.S.N.M. No. P969) and unfigured hypo-
types (U.S.N.M. No. P970) from the Annona chalk about 0.5 mile
north of the White Cliffs Post Office, Ark. Collected by W. H.
Deaderick.

Hypotypes of figures 4 and 5 (U.S.N.M. Nos. Pg71a-b) and un-
figured hypotypes (U.S.N.M. No. P972) from the Ozan sand (Upper
Cretaceous), in natural erosion on the east side of the Arkinda road,
5.0 miles northwest of Foreman, Little River County, Ark. Col-
lected by W. H. Deaderick.

Unfigured hypotypes (U.S.N.M. No. P973) from the Annona
chalk, about 0.5 mile west of the Brownstown—White Cliffs road, 1.7
miles south of Brownstown, Ark. Collected by W. H. Deaderick.

Unfigured hypotypes (U.S.N.M. No. P974) from the Annona
chalk (at the Ozan contact) on the east side of the road to White
Cliffs, 2.8 miles south of the Brownstown crossroads, Ark. Collected
by W. H. Deaderick.

Unfigured hypotypes (U.S.N.M. No. P975) from the Annona
chalk, on the east side of the road to Columbus, 1.0 mile south of
Yancey, Hempstead County, Ark. Collected by W. H. Deaderick.

Unfigured hypotypes (U.S.N.M. No. P976) from the Saratoga
chalk, on the south side of the road to Columbus, 2.0 miles east of
Saratoga, Howard County, Ark. Collected by W. H. Deaderick.

Unfigured hypotypes (U.S.N.M. No. P977) from the Upper
Cretaceous Nacatoch sand, on the north side of Highway 26, 0.5 mile
east of Big Decipher Creek, 5.0 miles west of the corner of 1oth and
Pine Streets in Arkadelphia, in SW34 sec. 27, T. 7 S., R. 20 W., Clark
County, Ark. Collected by W. H. Deaderick.

TRIPLASIA DEADERICKI Loeblich and Tappan, new species
Plate 6, figures 1-5

Test free, of medium size, elongate margins nearly parallel, early
portion planispirally coiled, later chambers uniserial and generally
triangular in section with somewhat rounded angles and slightly con-
cave sides, occasionally quadrate (fig. 4), and one specimen (fig. 1)
was found to have only two angles, and thus Flabellammina-like.
Occasional additional specimens are flattened in the early uniserial por-
tion, developing the third angle about one-half the distance from the
base, chambers numerous, low and broad in the early portion, later
ones higher, and final one or two chambers generally of smaller
diameter and rounded in section (figs. 1-3), and of approximately
equal height and breadth ; sutures obscure in the early portion, becom-

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

ing well marked and constricted in the later portion of the test, radial
in the coil, slightly arched in the rhomboid uniserial chambers and
horizontal between the later rounded chambers ; wall finely arenaceous,
with occasional coarser grains, usually rather smoothly finished;
aperture terminal, ovate to slitlike, slightly produced.

Length of holotype (fig. 3), 2.18 mm.; breadth, 0.78 mm. Length
of paratype of figure I, 1.77 mm.; breadth, 0.75 mm.; thickness
0.47 mm. Length of paratype of figure 2, I.90 mm.; breadth,
0.60 mm. Length of paratype of figure 4, 1.01 mm.; breadth, 0.60 mm.

Remarks.—This species resembles T. glenrosensis, new species, in
the rounded later chambers, large coil, and nearly parallel margins. It
differs in being narrower with sharper angles, more coarsely arena-
ceous, and with a roughened finish.

The species is named in honor of the late Dr. W. H. Deaderick, in
recognition of this work in assembling a superb collection of For-
aminifera from the Cretaceous strata of Arkansas.

Types and occurrence.—Holotype (fig. 3, U.S.N.M. No. Pg78),
paratypes of figures 2, 4, 5 (U.S.N.M. Nos. Pg7ga-c), and unfigured
paratypes (U.S.N.M. No. Po80) from the Upper Cretaceous Annona
chalk, at the Ozan contact, on the east side of the road to White
Cliffs, 2.8 miles south of the Brownstown crossroads, Ark. Collected
by W. H. Deaderick.

Paratype of figure 1 (U.S.N.M. No. Po81) and unfigured para-
types (U.S.N.M. No. P982) from the Annona chalk, about 0.25 mile
north of the White Cliffs Post Office, Ark. Collected by W. H.
Deaderick.

TRIPLASIA PLUMMERAE Loeblich and Tappan, new species
Plate 6, figures 14-19b

Test free, large, elongate, triangular or occasionally quadrate in
section, angles rounded, sides moderately excavated, slightly flaring to
subparallel-sided, uniserial throughout, base occasionally slightly
curved ; chambers numerous, low, increasing very gradually in height
as added, each succeeding chamber overlapping the preceding, final
chamber nearly as high as broad, chambers extending sharply down-
ward at the angles ; sutures fairly distinct, slightly depressed, especially
in the later portion of the test, strongly arched on the faces of the test,
recurved at the angles; wall arenaceous, with grains of medium size
in a ground mass of finer material and occasionally incorporating tests
of smaller species of Foraminifera; aperture terminal, a narrow slit,
occasionally slightly produced on a short neck.

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 37

Length of holotype (fig. 19), 5.46 mm.; breadth, 1.74 mm. Length
of quadrate paratype of figure 17, 4.34 mm.; breadth, 1.46 mm.
Other paratypes range from 2.21 to 5.41 mm. in length and from
0.86 to 2.29 mm. in breadth.

Remarks.—This species is somewhat variable in character, two
specimens from the Saratoga chalk of Arkansas showing a compressed
Flabellammina-like early portion, although no distinct coil is present,
and the third angle is only developed in the latter half of the test.
The majority of specimens are regularly triangular in section, but
four of the paratypes are quadrate, two of these from the Taylor
marl of Texas being figured, the other two occurring in the Saratoga
chalk of Arkansas. As 156 specimens of this species were examined,
approximately 2.6 percent of the specimens are quadrate and about 1.3
percent are flattened, lacking the third angle in the early stage.

Although a few specimens show a slight curve at the base (fig. 14)
there is no definite coil.

The present species differs from Triplasia murchisoni Reuss in
being less flaring, with less-excavated sides and more-rounded final
chambers. T. murchisoni Reuss is approximately three-fifths as large
as the present species and has a rounded rather than elongate aperture.

Triplasia taylorensis (Cushman and Waters), which occurs with
this species, has a more flaring test with a definite coil at the base and
more deeply excavated sides, and tends to be more coarsely arenaceous,
incorporating larger fragments in the wall.

Types and occurrence-—Holotype (fig. 19, U.S.N.M. No. P983),
paratypes of figures 14-18 (U.S.N.M. Nos. Po8q4a-e), and unfigured
paratypes (U.S.N.M. No. P985) all from the greenish-gray marls
of the upper Taylor, on the right bank of Onion Creek, just down-
stream from the bridge at Moore and Berry’s Crossing, 8.5 miles
southeast of the State Capitol building in Austin, Travis County, Tex.
Collected by Helen Tappan Loeblich and Alfred R. Loeblich, Jr.

Paratype (U.S.N.M. No. P986) from the Saratoga chalk, on U. S.
Highway 51, 10.5 miles southwest of Wright’s store at the junction
of highways 26 and 51, 3.1 miles northeast of Okalona, on the
northeast side of the road, Clark County, Ark. W. H. Deaderick
collection in the U. S. National Museum.

Paratypes (U.S.N.M. No. P987) from the Saratoga chalk, 2.0 miles
east of Saratoga, on the south side of the road to Columbus, Howard
County, Ark. W. H. Deaderick collection in the U. S. National
Museum.

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

TRIPLASIA ROMERI (Reuss)
Plate 5, figures 21a-22b

Rhabdogonium rémeri Reuss, 1860, Sitzb, Akad. Wiss. Wien, math.-naturw. K1.,
vol. 40, p. 201, pl. 6, figs. 7a-c.
Ammobaculites roemeri (Reuss) FRANKE, 1928, Abh. preuss. geol. Landesanst.,
Berlin, n.s., No. 111, p. 166, pl. 15, fig. 8.
Original description (translated from German) :

Test elongate, 1.97 mm. in length by 0.643 mm. in breadth, occasionally some-
what curved, of almost equal breadth for the entire length, at the upper end
short and bluntly acuminate, sharply rounded below or tapering to a blunt end,
sharply triangular, with nearly equal sides. Three to six 3-sided chambers, uni-
form, with shallow, very weakly curved sutures and sharp angles; only the
last chamber shows somewhat convex faces and rounded angles. They have the
form of a 3-sided pyramid, whose rounded apex bears the round simple aperture.
Surface of the test rough. Very rare.

Remarks.—This species somewhat resembles T. deadericki, new spe-
cies, from the Annona chalk of Arkansas, in general shape and the
relatively high and somewhat rounded later chambers. It differs in hav-
ing more distinct angles and somewhat convex sides instead of the
slightly concave sides and broadly rounded angles of the American
species.

Types and occurrence.—Originally described from the upper Seno-
nian (Mucronatenkreide) marl at Hilgenberg bei Hamm, Westphalia,
Germany. It was also recorded from the upper Senonian of West-
phalia by Franke.

Hypotype of figure 22 (U.S.N.M. No. P988) from the upper
Senonian, Dasbeck bei Hamm, Germany. Collected by A. Franke
and given to the U. S. National Museum by C. I. Alexander.

TRIPLASIA RUGOSISSIMA (Alexander and Smith)
Plate 7, figures 6a-9

Frankeina rugosissima ALEXANDER and SMITH, 1932, Journ. Paleontol., vol. 6,
No. 4, p. 311, pl. 47, figs. 12-13—-CuUSHMAN, 1946, U. S. Geol. Surv. Prof.
Pap. 206, p. 25, pl. 4, figs. 13-14.

Test free, large, flaring from the base, later with nearly parallel
sides, with a small planispiral coil at the base which is apparently ob-
solete in some specimens (fig. 9), the major portion of the test uni-
serial and triangular in section, occasionally quadrate (fig. 6), angles
sharp, sides moderately excavated; chambers broad, gently arched ;
sutures obscure, but discernible on some specimens; wall coarsely
arenaceous, surface rough with many fragments of shells, fish bones,

NO. I5 TRIPLASIA REUSS—-LOEBLICH AND TAPPAN 39

occasional ostracod carapaces, glauconite and other dark mineral
grains mixed with the large quartz grains, in a ground mass of finer
material; aperture terminal, elongate.

Length of holotype (fig. 7), 3.75 mm.; breadth, 1.85 mm. Length
of quadrate hypotype of figure 6, 2.47 mm. ; breadth, 1.12 mm. Length
of hypotype of figure 9, 3.59 mm. ; breadth, 1.61 mm. Other hypotypes
range in length from 0.96 to 3.85 mm.

Remarks.—Alexander and Smith stated that this species occurred
only in the lower middle Taylor and was geographically restricted to
southern Texas. We have specimens that are identical in all charac-
teristics from the Saratoga and Annona chalks of Arkansas. This is
one of the largest and most coarsely agglutinated species of this genus.

Types and occurrence——Holotype (fig. 7, U.S.N.M. No. Pgg93),
paratypes figured by Alexander and Smith (1932, pl. 47, fig. 12;
U.S.N.M. No. P994), and unfigured paratypes (Cushman coll. No.
17851) all from about 150 feet above the base of the Upper Cre-
taceous Taylor clay, in a gully north of the Austin-Manor highway,
near the east end of the long bridge over Walnut Creek, 6.0 miles
northeast of Austin, Travis County, Tex. Collected by C. I. Alexander.

Unfigured hypotypes (U.S.N.M. No. P995) from the Upper Cre-
taceous Saratoga chalk from the top of Devil’s Backbone, Howard
County, Ark. Collected by W. H. Deaderick.

Unfigured hypotypes (U.S.N.M. No. P996) from the Annona
chalk at the Ozan contact, 2.8 miles south of the Brownstown cross-
roads on the east side of the road to White Cliffs, Ark. Collected by
W. H. Deaderick.

Hypotypes of figures 6, 8, 9 (U.S.N.M. Nos. Pgg7a-c) and un-
figured hypotypes (U.S.N.M. No. Pg998) from a natural erosion of
the Annona chalk about 0.5 mile north of the White Cliffs Post Office,
Ark. Collected by W. H. Deaderick.

TRIPLASIA TAYLORENSIS (Cushman and Waters)
Plate 6, figures 6-13

Frankeina taylorensis CUSHMAN and WATERS, 1929, Contr. Cushman Lab.
Foram. Res., vol. 5, p. 63, pl. 10, figs. 3a-b—ALEXANDER and SMITH, 1932,
Journ. Paleontol., vol. 6, No. 4, p. 310, pl. 47, figs. 7, 9 -CUSHMAN, 1946,
U4S. Geol. Surv. Prof. Pap. 206, p. 25, pl. 5, figs. 1-2.

Test free, large, elongate, somewhat flaring, planispiral coil very
small or obsolete, uniserial portion generally triangular, occasionally
quadrate (about 0.9 percent of the specimens observed), sides mod-
erately excavated, angles acute; chambers numerous, low and broad,

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

increasing very little in height but increasing somewhat in breadth
in the early portion so that the test is flaring at first, but later has
nearly parallel sides, final chamber somewhat higher than the pre-
ceding ; sutures distinct, slightly depressed, gently arched on the sides,
recurved at the angles; wall arenaceous, of uniformly sized, moder-
ately coarse grains of quartz with a scattering of dark mineral grains,
surface even, but very granular in appearance; aperture terminal,
slitlike, not produced.

Length of holotype (fig. 8), 2.13 mm.; breadth, 0.96 mm. Length
of hypotype of figure 6, 3.61 mm.; breadth, 1.56 mm.; thickness,
1.51 mm. Length of hypotype of figure 7, 3.46 mm.; breadth,
1.40mm, Length of hypotype of figure 9, 1.51 mm. ; breadth, 0.62 mm.
Length of hypotype of figure 10, 3.17 mm. Length of hypotype of
figure II, 3.77 mm.

Remarks.—Among the holotype, three paratypes, and 437 hypo-
types, there are four quadrate specimens, the remainder being tri-
angular. The species is characterized by its large size, acute angles,
excavated faces, low chambers and reduced early coil.

Types and occurrence-—Holotype (fig. 8, Cushman coll. No. 12033)
and paratype (Cushman coll. No. 12034) from the Upper Cretaceous
Taylor marl at 360 feet depth, and paratypes (Cushman coll. No.
12035) from the Taylor marl at a depth of 365 feet, all from the Sun
Oil Company, Martindale D 10, in Caldwell County, Tex.

Hypotypes figured by Alexander and Smith, 1932 (U.S.N.M. No.
Pg99) from the upper Taylor marls, in the bank of a small stream
200 yards south of the bridge on Pierce’s Lane, 1.5 miles southeast of
Del Valle, in Travis County, Tex. Collected by C. I. Alexander.

Hypotypes of figures 6, 7, 9-13 (U.S.N.M. Nos. Proooa-g) and
unfigured hypotypes (U.S.N.M. No. Proor) from the upper Taylor
marl, in the right bank of Onion Creek, just downstream from the
bridge at Moore and Berry’s Crossing, 8.5 miles southeast of the State
Capitol building, in Austin, Travis County, Tex. Collected by Helen
Tappan Loeblich and Alfred R. Loeblich, Jr.

Hypotypes (U.S.N.M. No. Pioo2) from the Upper Cretaceous
Annona chalk, about 0.5 mile north of the White Cliffs Post Office,
Ark. Collected by W. H. Deaderick. ®

Hypotypes (U.S.N.M. No. P1003) from the Annona chalk, about
0.5 mile west of the road between Brownstown and White Cliffs, and
about 1.7 miles south of Brownstown, Ark. Collected by W. H.
Deaderick.

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN AI

Hypotypes (U.S.N.M. No. Pioo4) from the Annona chalk, 1.0
mile south of Yancey, on the east side of the road to Columbus, in
Hempstead County, Ark. Collected by W. H. Deaderick.

TRIPLASIA SARATOGENSIS Loeblich and Tappan, new species
Plate 7, figures 1oa-11b

Test free, narrow and elongate, of nearly uniform breadth for most
of its length, triangular to quadrate in section, sides nearly flat to
slightly concave, angles broadly rounded ; chambers usually indistinct,
with a coil of about three chambers at the base, followed by uniserial
chambers, final chamber slightly produced to the aperture ; sutures in-
distinct, occasionally visible and gently arched on the faces of the
test; wall arenaceous, of medium to coarse grains in a finer ground
mass, surface moderately rough in appearance; aperture terminal,
ovate.

Length of holotype, 2.47 mm.; breadth, 0.91 mm. Length of quad-
rate paratype, 2.76 mm.; breadth, 0.75 mm. Other paratypes range
from 1.01 to 3.25 mm. in length.

Remarks.—This species differs from Triplasia cushmani (Alexan-
der and Smith) in being longer and comparatively more narrow, and
in having a less well-defined coil and less-depressed sutures. The
sides are also flat, while those of T. cushmani are excavated and the
angles sharper. Triplasia rugosissima (Alexander and Smith) has a
larger, much broader and more flaring test. T. deadericki, new species,
has a smaller test, with higher chambers and much more constricted
sutures.

Types and occurrence.—Holotype, figure 10 (U.S.N.M. No.
P1005), quadrate paratype of figure 11 (U.S.N.M. No. P1006), and
unfigured paratypes (U.S.N.M. No. Pioo7) from the Saratoga
chalk, at the top of Devil’s Backbone, in Howard County, Ark. Col-
lected by W. H. Deaderick.

Unfigured paratypes (U.S.N.M. No. P1008) from the Saratoga
chalk, in natural erosion just west of Saratoga, Howard County, Ark.
Collected by W. H. Deaderick.

Unfigured paratypes (U.S.N.M. No. Pioog) from the Saratoga
chalk, on the north side of the road, at the top of a high hill, 2.0 miles
east of the junction of Highways 73 and 55 in Saratoga, Howard
County, Ark. Collected by W. H. Deaderick.

Unfigured paratypes (U.S.N.M. No. Proro) from the Saratoga

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

chalk, on the east side of the road, just north of Saratoga, Howard
County, Ark. Collected by W. H. Deaderick.

Unfigured paratypes (U.S.N.M. No. Pio11) from the Saratoga
chalk, on the south side of the road to Columbus, 2.0 miles east of
Saratoga, Howard County, Ark. Collected by W. H. Deaderick.

Unfigured paratypes (U.S.N.M. No. Pio12) from the Saratoga
chalk, on the east side of Highway 29, 1.8 miles south of Blevins,
13.5 miles north of Hope, Hempstead County, Ark. Collected by
W. H. Deaderick.

Unfigured paratypes (U.S.N.M. No. Pro13) from the Saratoga
chalk, on the east side of Highway 4, on the road to Ozan, 3.1 miles
north of Washington, Hempstead County, Ark. Collected by W. H.
Deaderick.

TRIPLASIA species
Plate 7, figures 12a, b

Frankeina sp. IsRAELSKY, 1951, U. S. Geol. Surv. Prof. Pap. 240-A, p. 12, pl. 3,
figs. 5-8.

Test free, of medium size, with a large planispiral coil of about five
chambers, followed by a few uniserial chambers with nearly parallel
margins and triangular in section, the two angles in the plane of
coiling more prominent and the third angle less elevated, sides flat,
angles rounded; chambers low and broad; sutures radiate in the
coil, slightly arched on the faces of the triangular portion, distinct
and slightly depressed ; wall coarsely arenaceous, with occasional dark
mineral grains, surface rough in appearance; aperture terminal,
elongate.

Length of figured specimen, 1.69 mm.; breadth of coil, 0.73 mm.;
greatest breadth of triserial portion, 0.68 mm.

Remarks.—This species resembles T. glenrosensis, new species, in
the large coil and less prominent third angle. It differs in having lower
chambers, more acute angles and more coarsely arenaceous and more
roughly finished wall. The species is represented by a single speci-
men, and because a single specimen often is not representative of
the complete species of this variable genus, this form has not been
named.

Types and occurrence.—Figured specimen (U.S.N.M. No. 560500)
from the Lodo formation (Paleocene and Eocene in age), greenish-
gray, silty calcareous claystone, 960 feet below the Middle Eocene
Domengine formation, sec. 29, T. 15 S., R. 12 E., northwest Tumey
Hills, Fresno County, Calif. Collected by M. C. Israelsky.

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 43

TRIPLASIA FUNDIBULARIS (Harris and Jobe)
Plate 7, figures 13a-14b

Frankeina fundibularis Harris and Jose, 1951, Microfauna of basal Midway
outcrops near Hope, Arkansas, p. 7, pl. 1, figs. 8a-c.

Test free, of medium size for the genus, elongate, with a large
early planispiral portion and later triangular portion, sides moderately
to deeply excavated, angles acute; chambers increasing in size as
added, low and broad, slightly arched on the faces, extending down-
ward at the angles; sutures distinct, slightly depressed; wall very
coarsely arenaceous, incorporating smaller Foraminifera and organic
fragments with the sand and other mineral grains, surface rough;
aperture terminal, rounded.

Length of hypotype of figure 13, 1.87 mm.; breadth, 0.99 mm.
Length of hypotype of figure 14, 1.27 mm.; breadth, 0.62 mm. Other
hypotypes range from 1.09 to 1.98 mm. in length.

Remarks.—Only a few specimens (seven in all) were available to
the writers, and all of these were triangular. It is probable that a
very large suite of specimens would show the quadrate or flattened
modifications. This species differs from Triplasia species, here figured
from the Paleocene Lodo formation of California, in being more
flaring and more sharply angled, and in having a somewhat smaller
planispiral portion and more highly arched sutures.

Types and occurrence —The holotype (deposited in the University of
Oklahoma collection), figured hypotypes (U.S.N.M. Nos. Piorga, b)
and unfigured hypotypes (U.S.N.M. No. Pior5) are all from the
Paleocene Midway formation, 1.25 miles south of Terre Rouge Creek
on State Highway 29, 5 miles north of Hope, sec. 9, T. 12 S., R. 24 W.,
Hempstead County, Ark. Collected by Mrs. Billye Jobe.

TRIPLASIA ANDRAEI (Liebus)
Text figure II

Haplophragmium andraei Liesus, 1911, Sitzb. Akad. Wiss. Wien, math.-naturw.
K1., vol. 120, pt. I, p. 940, text figs. 5a-c.

Original description (translation from German) :

The test is over 1.5 mm. long, and very coarsely agglutinated, so that the
chamber arrangement is almost completely obscured from the exterior. The
center of the test’s breadth is very highly elevated, in certain places so strongly
inflated that it has the appearance of padding, extending from about one-eighth
of the distance from the base of the test almost six-eighths of the entire length.
The ends of this swelling slope very sharply down to the remaining upper sur-
face of the test. At the end of the test is found a short projection, bearing a
somewhat broad and distended aperture.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

First in the development one observes 5 chambers spirally arranged in the
older portion of the test, to which the remaining uniserial ones are then joined;
concerning the relationship to Haplophragmiwm there can be no doubt. None
of the previously known forms of this genus show a 4-sided cross section
caused by the inflated swelling and the flattening of the surrounding area.

The form also varies somewhat, however, and these variations do not show
the inflated elevation of the broad side to such a height as the example here

Fic. 11.—Triplasia andraeit (Liebus): a, Side view of holotype. b, Top
view, showing aperture and quadrate section. c, Sectional view, showing early
coil followed by uniserial chambers. (After Liebus. )

figured. These specimens tend toward the form that Andrae recorded under the
name Haplophragmium humboldti Reuss var. latum, n. var. A comparison with
the former occurrence of H. humboldti Reuss shows that an eventual gradation
of Andrae’s form into the true H. humboldti Reuss does not exist. The typical
H. humboldti Reuss is somewhat compressed in its older portion but has, how-
ever, without exception, a round cross section in the younger uniserial portion,
whereas in the present form the younger part still earlier demonstrates a
strong beveling of the test.

For the present, as long as the material is still rare—there are 4 specimens
observed—I consider this species as identical with the form that Andrae de-

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 45

scribed as H. humboldti Reuss var. latum although differing from the true
H. humboldti Reuss and propose as a new designation Haplophragmium andraci.

For these Haplophragmiums that at the end are composed of uniserially ar-
ranged chambers, Cushman has recently proposed the name Ammobaculites,
whereas he placed the completely enrolled forms in Haplophragmoides. Accord-
ing to this definition the preceding would be known as Ammobaculites andraci
n. Sp.

Remarks.—The nomenclature of this species is quite complicated.
As quoted above, Liebus proposed this specific name, figuring a defi-
nite quadrate Triplasia, but in his description stating that it was identi-
cal to H. humboldti: Reuss var. latum Andrae.

On this basis, Ellis and Messina (Catalogue of Foraminifera) refer
to the present species as Haplophragmium andraei Liebus, 1911, nom.
nov. If the species had been so described, the type of Andrae’s species
would have thus become the type for the new name and Liebus
would have had no basis for proposing a new specific name, but should
have elevated Andrae’s variety latum to specific rank. Liebus’s name
thus would be a synonym for Haplophragmium latum (Andrae).

However, Liebus erroneously considered Andrae’s variety to be
conspecific, as the latter is not angular in section, but ovate, and as it
has horizontal sutures in this compressed uniserial portion it should
thus be placed in Ammobaculites.

Furthermore, Liebus described his species under the heading n. sp.
and not nom. nov., and his specific description referred to the quad-
rate specimen he figured (shown here in text figure 11) and not to
the specimens of Andrae. Therefore, it was apparently his intent to
describe a new species for his quadrate specimen, and in fact in the
last paragraph of his discussion he stated that it might belong to
Ammobaculites and would be known then as Ammobaculites andraei,
new species. Therefore his name is here recognized as a valid specific
name for this quadrate Triplasia.

Occurrence.—Middle Eocene beds in a well at the village of Kolar-
ine, northern Dalmatia.

TRIPLASIA HUNGARICA (Majzon)
Plate 8, figures Ia-c

Centenarina hungarica Majzon, 1948, Foldtani Kozlony, vol. 78, p. 24, figs.
ite MEL op

The following is a free translation from the Hungarian:

In the first stage of development appears a flat planispiral portion of four to
five chambers and later there is a uniserial development of arched chambers

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

which are rectangular in cross section. Their number cannot be determined
exactly because of the rough sandy material of the test. They form approxi-
mately four concave rectangular prisms that are somewhat produced terminally
where the aperture is slit-shaped. The sutures in the coiled portion are radial
and in the 4-sided prismatic portion they are somewhat arched.

Length, 1.6 mm.; width, 0.65 mm.

RRemarks.—This species was described as the type species of the
genus Centenarina Majzon, but the latter name is a junior synonym of
Triplasia and is here suppressed. T. hungarica resembles quadrangular
forms of T. variabilis (see pl. 8, figs. 5, 9) in shape, excavation of
sides, and slitlike aperture, but is only one-half as large and has a more
inflated early portion.

Occurrence.—( Translation) “It is found in the Farkasreti cemetery
in Budapest, in the fourth foraminiferal horizon of the Rupelian
(Oligocene) strata.”

TRIPLASIA TRIGONA (Andrae)

Haplophragmium humboldti (Reuss) var. trigona ANDRAE, 1800, Mitt. geol.
Landesanst., Alsace-Lorraine, vol. 3, No. 1, p. 116.

The original description (translation from German) states, “This
variety differs from the typical form in that the last chambers be-
come triangular, as are many Clavulinas and Tritaxias.”

Remarks.—As Spirolina humboldti Reuss is an Ammobaculites,
and the triangular section of this “variety” places the present form in
Triplasia, Andrae’s varietal name is here raised to specific rank. Un-
fortunately this species has never been figured, and no specimens were
available to the writers.

Types and occurrence —Middle Oligocene, Septarienthon, Lobsann
and Sulz unterm Wald, Unter-Elsass. The types are deposited in the
collection of the Geologische Landesanstalt of Alsace-Lorraine, Strass-
bourg, France.

TRIPLASIA MARWICK! Loeblich and Tappan, new species
Plate 8, figures 2-4b

Test free, large, elongate, planispiral coil not evident, early portion
flaring, later triangular to quadrate in section, sides flat to slightly
excavated, angles rounded; chambers low in the early portion, in-
creasing somewhat in height as added; sutures obscure, gently arched
at the center of the faces, recurved at the angles; wall arenaceous,
with occasional very large grains incorporated in the main body of

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 47

finer-grained material, surface rather smoothly finished; aperture a
terminal slit.

Length of holotype (fig. 4), 3.59 mm.; breadth, 1.53 mm. Length
of small paratype of figure 2, 1.69 mm. ; breadth, 1.33 mm. Length of
paratype of figure 3, 3.04 mm.; breadth, 1.51 mm.

Remarks.—This species is named in honor of Dr. J. Marwick,
Director of the New Zealand Geological Survey, in recognition of
his work on the Upper Cretaceous and Tertiary of New Zealand.

It differs from T. rugosissima (Alexander and Smith) in being
broader with less concave sides and lower chambers, and in lacking
the slight neck of the Cretaceous species.

Types and occurrence-——Holotype (U.S.N.M. No. Pio16) and
unfigured paratype (U.S.N.M. No. Pro17) from the Miocene of
Kawhia, New Zealand. Paratypes of figures 2 and 3 (U.S.N.M. Nos.
Pio18a, b) and unfigured paratype (U.S.N.M. No. Proig) from the
Miocene of Motatura, New Zealand.

TRIPLASIA MINUTA (Reuss)
Plate 7, figures 16a-17b

Rhabdogonium minutum Reuss, 1867, Sitzb. Akad. Wiss. Wien, math.-naturw.
K1., vol. 55, pt. 1, p. 85, pl. 5, figs. 4a-5b.

not Rhabdogonium minutum Reuss, Brapy, 1884, Rep. Voy. Challenger, Zool.,
vol. 9, p. 526, pl. 67, figs. 4-6.

Original description (translation from German) :

In this species with very fragile margins two very different principal forms
may be distinguished. One is larger as a rule, inverted-ovate in side view,
wider above than below, quite often considerably elongated, often irregularly
twisted. The surface of the shell is rougher above.

Other tests are, on the other hand, often more regularly oval, tapering at
both ends, generally regularly formed and with even margins. In all cases,
however, the three lateral angles are thin, flangelike, the sides concave. The
development of the uniserial chambers lying in a straight line one on another
is only sporadically indicated by the very slight, transverse, gently curved fur-
rows. The terminal round aperture is small, often indistinct. From this, in the
individuals which are broader above, extend three fine, short furrows which lie
upon the upper thicker portion of the angles.

In appearance this species, which is found to be common in Salzthone, is very
close to Tritaxia tricarinata Reuss from the Upper Cretaceous.

Remarks.—This species has not been seen by the writers, but is
here considered to belong to Triplasia, on the basis of Reuss’s figures
and descriptions and because he referred it to Rhabdogonium (of
which he was the author, proposing it to supersede Triplasia when

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

he found some specimens to be quadrate). It is the only Miocene
species of this genus previously described.
Occurrence.—Miocene, Salzthone, at Wieliczka, Galizien, Poland.

TRIPLASIA WRIGHTI (De Amicis)
Plate 7, figures 18a-b

Haplophragmium wrighti De Amicts, 1895, Natur. Siciliano, Palermo, Ann. 14
(1894-95), Nos. 4-5, p. 58, pl. 1, figs. 12a-b.

This species was described as having an initial spire, followed by
uniserial triangular chambers. The coil is comparatively large, the
sides of the triangular portion flat to slightly convex, and the angles
unusually sharp, the uniserial portion has parallel margins and strongly
constricted sutures. The aperture is terminal and rounded. The type
is 2.8 mm. in length and only a single specimen was found.

Remarks.—This is the only Pliocene species recorded. It somewhat
resembles T. romeri in the large coil, and rather high, slightly in-
flated uniserial chambers, with parallel margins. It differs in having
much sharper angles, and is about one-third larger.

Type and occurrence.—Type specimen from the Lower Pliocene of
Bonfornello, presso Termini-Imerese, Sicilia, Italy.

TRIPLASIA VARIABILIS (Brady)
Plate 8, figures 5-9b

Verneuilina variabilis Brapy, 1884, Rep. Voy. Challenger, Zool., vol. 9, p. 385,
pl. 47, figs. 21-24.

not Ammobaculites variabilis Brady, FRANKE, 1928, Abh. preuss. geol. Lan-
desanst., Berlin, n.s., No. 111, p. 166, pl. 15, figs. 6a-b.

Test large, small specimens or early portion of larger ones flaring,
later portion nearly parallel-sided, triangular to quadrangular in sec-
tion, and a few specimens may lack the third angle and be flattened,
angles acute, sides moderately excavated, chambers numerous, broad
and low, gently arched; sutures obscure to well marked and slightly
depressed; wall coarsely arenaceous, surface rough; aperture a ter-
minal slit, only very slightly produced, if at all.

Brady gave the length as 3 mm. The hypotype here shown in figure
5 is 2.86 mm. in length and 1.43 mm. in breadth. Length of hypotype
of figure 7, 3.80 mm., and breadth, 1.85 mm. Length of hypotype of
figure 9, 3.77 mm.; breadth, 2.11 mm.

Remarks.—Brady described this species as typically triangular, but
stated that it was polymorphic: “Of the numberless modifications of

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 49

the typical structure, the most common are . . . those . . . in which
the test is more or less quadrate and . . . in which it loses to some
extent its ternate character in the other direction and presents during
part of its growth only two marginal edges. No specific or even
varietal distinction can be drawn between these forms; and...
every intermediate condition is common.” Brady thus figured tri-
angular and quadrate specimens as well as the flattened forms re-
sembling Flabellammina. A parallel development is shown in the
Lower Cretaceous Triplasia incerta (Alexander and Smith), some
of which show a Flabellammina stage of varying length before the
third angle appears, and in the Lower Jurassic Triplasia kingakensis
which also has flattened, triangular, and quadrate forms.

Franke (1928, p. 166) referred to this species Upper Cretaceous
specimens (Senonian) but placed the species in the genus Ammobac-
ulites. His specimens seem closer to Triplasia murchisoni Reuss,
however.

Majzon (1948, p. 24) in describing a 4-sided species as the type for
his genus Centenarina, stated in footnotes (free translation from the
Hungarian), “In my opinion, Brady’s Verneuilina variabilis is Franke-
ina,’ and “One of Brady’s figures of the above mentioned V. vari-
abilis may possibly belong to Centenarina.”

Thus these specimens figured by Brady have been variously re-
ferred to the genera Verneuilina (by Brady), Ammobaculites (Franke,
1928, p. 166), Triplasia (Bartenstein and Brand, 1937, p. 185), Frank-
eina (Majzon, 1948, p. 24), and Centenarina (Majzon, 1948, p. 24).

Types and occurrence.—Brady’s types from the coral sands of
Kandavu, Fiji Islands, at 210 fathoms, are in the British Museum at
South Kensington and in the Brady collection in the Museum of
Comparative Anatomy at Cambridge, England. Figured hypotypes
(U.S.N.M. Nos. Pio20a-e) and unfigured hypotypes (U.S.N.M.
No. Pro21) from Challenger station 174-D, from coral sands at
210 fathoms, off Kandavu, Fiji Islands, at lat. 19°6’ S., long. 178°14’
20” E., collected March 8, 1874.

REFERENCES

ALExANDER, C. I., and Smit, J. P.
1932. Foraminifera of the genera Flabellammina and Frankeina from the
Cretaceous of Texas. Journ. Paleontol., vol. 6, No. 4, pp. 299-311,
pls. 45-47.
ANpRAE, A.
1890. Weitere Beitrage zur Kenntniss des Oligocans im Elsass. Mitt. geol.
Landesanst., Alsace-Lorraine, Strassbourg, vol. 3, No. I, pp. 105-122,
pls. 5-6.

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

BarTENSTEIN, H., and Branp, E.

1937. Mikro-palaontologische Untersuchungen zur Stratigraphie des nord-
westdeutschen Lias und Doggers. Abh. Senckenberg. naturf. Ges.,
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1949. New genera of Foraminifera from the Lower Cretaceous of Germany
and England. Journ. Paleontol., vol. 23, No. 6, pp. 669-672, 10 figs.

1951. Mikropalaontologische Untersuchungen zur Stratigraphie des nord-
westdeutschen Valendis. Abh. Senckenberg. naturf. Ges., No. 485,
Festschrift Rudolf Richter, pp. 239-336, pls. 1-25.

BEtsseEL, I.

1891. Die Foraminiferen der Aachener Kreide. Abh. preuss. geol. Lan-

desanst., Berlin, n.s., No. 3, pp. 1-78, pls. 1-16.
Brapy, H. B.

1884. Report on the Foraminifera dredged by H. M. S. Challenger during the
years 1873-1876. Rep. Voy. Challenger, Zool., vol. 9, pp. 1-814,
pls. I-115.

CusHMAN, J. A.

1933. Foraminifera, their classification and economic use. 2d ed. Cushman
Lab. Foram. Res. Spec. Publ. No. 4, pp. 1-340.

1946. Upper Cretaceous Foraminifera of the Gulf coastal region of the
United States and adjacent areas. U. S. Geol. Surv. Prof. Pap. 206,
pp. 1-241, pls. 1-66.

1948. Foraminifera, their classification and economic use. 4th ed., pp. 1-605,
pls. 1-55. Harvard Univ. Press.

CusHMAN, J. A., and ALEXANDER, C. I.

1929. Frankeina, a new genus of arenaceous Foraminifera. Contr. Cushman

Lab. Foram. Res., vol. 5, pp. 61-62, pl. ro.
CusHMAN, J. A., and Ponton, G. M.

1932. An Eocene foraminiferal fauna of Wilcox age from Alabama. Contr.

Cushman Lab. Foram. Res., vol. 8, pp. 51-72, pls. 7-9.
Cusuman, J. A., and Waters, J. A.

1929. Some arenaceous Foraminifera from the Taylor marl of Texas.

Contr. Cushman Lab. Foram. Res., vol. 5, pp. 63-66, pl. 10.
De Amicis, G. A.

1895. I Foraminiferi del pliocene inferiore di Bonfornello. Natur. Siciliano,

Palermo, Ann. 14, Nos. 4-5, pp. 51-74, 1 pl.
Eccrer, J. G.

1899. Foraminiferen und Ostrakoden aus den Kreidemergel der Ober-
bayerischen Alpen. Abh. bayer. Akad. Wiss., Mtinchen, Kl. 2,
vol. 21, pt. I, pp. 1-230, pls. 1-27.

EICHENBERG, W.

1934. Die Erforschung der Mikro-organismen, insbesondere der Foramini-
feren der norddeutschen Erdoélfelder. Teil 1. Die Foraminiferen
der Unterkreide. Folge 3. Foraminiferen aus dem Hauterive von
Wenden am Mittellandkanal. Niedersachs. geol. Ver. Hannover,
Jahrb. 26, pp. 150-196, pls. 10-17.

FRANKE, A.

1928. Die Foraminiferen der Oberen Kreide Nord—und Mitteldeutschlands.
Abh. preuss. geol. Landesanst., Berlin, n.s., No. III, pp. 1-207,
pls. 1-18.

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 51

Harris, R. W., and Joss, B. I.
1951. Microfauna of basal Midway outcrops near Hope, Arkansas, pp. I-113,
pls. 1-14. Norman, Okla.
IsraELsky, M. C.
1951. Foraminifera of the Lodo formation, central California. U. S. Geol.
Surv. Prof. Pap. 240-A, pp. 1-29, pls. 1-11.
KeEtuer, B. M.
1946. The Foraminifera of the Upper Cretaceous deposits in the Sotchi
region. Bull. Soc. Natur., Moscow, n.s., vol. 51 (Sect. Géol., vol.
21), No. 3, pp. 83-108.
Ligesus, A.
1911. Die Foraminiferenfauna der mitteleocanen Mergel von Norddalmatien.
Sitzb. Akad. Wiss. Wien, math.-naturw. K1., vol. 120, pt. 1, pp. 865-
950, 3 pls.
LoesticH, A. R., Jr., and TAppAN, HELEN.
1950. North American Jurassic Foraminifera II: Characteristic western
interior Callovian species. Journ. Washington Acad. Sci., vol. 40,
No. 1, pp. 5-19, 1 pl.
ILOvAG), 18, Ley, fie
1944. Biostratigraphic relations of some north Texas Trinity and Fred-
ericksburg (Comanchean) Foraminifera. Amer. Midl. Nat., vol. 31,
No. 3, pp. 513-582, 5 pls.
Majzon, L.
1948. Centenarina, n. gen., and Cassidulina vitdlisi, n. sp., from the Lower
Rupelian strata at Budai. FGldtani Kozlony, vol. 78, pp. 22-25.
Marts, P.
1941. Les foraminiféres de la Craie a Belemnitella mucronata du Bassin de
Paris. Mém. Mus. Nat. Hist. Natur., Paris, n.s., vol. 12, No. 1,
pp. 1-296, pls. 1-37.
MYJATLIUK, E.
1939. The Foraminifera from the Upper Jurassic and Lower Cretaceous
deposits of the middle Volga region and Obschiy Syrt. Neftianyi
Geol. Razved. Inst. (Trans. Geol. Oil Inst.), Trudy, ser. A, vol.
120, pp. 3-76, pls. 1-4.
Reuss, A. E.
1854. Beitrige zur Charakteristik der Kreideschichten in den Ostalpen,
besonders im Gosauthale und am Wolfgangsee. Denkschr. Akad.
Wiss. Wien, math.-naturw. K1., vol. 7, pp. 1-156, pls. I-31.
1860. Die Foraminiferen der Westphalischen Kreideformation. Sitzb. Akad.
Wiss. Wien, math.-naturw. K1., vol. 40, pp. 147-238.
1867. Die fossil Fauna der Steinsalzablagerung von Wieliczka in Galizien.
Sitzb. Akad. Wiss. Wien, math.-naturw. K1., vol. 55, pt. I, pp.
17-182, pls. 1-8.
Steap, F. L.
1951. Foraminifera of the Glen Rose formation (Lower Cretaceous) of
central Texas. Texas Journ. Sci., vol. 3, No. 4, pp. 577-605, pls. 1-3.
Tappan, H.
1943. Foraminifera from the Duck Creek formation of Oklahoma and
Texas. Journ. Paleontol., vol. 17, No. 5, pp. 476-517, pls. 77-83.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

TEN Daw, A. ;
1946. Arenaceous Foraminifera and Lagenidae from the Neocomian

(Lower Cretaceous) of the Netherlands. Journ. Paleontol., vol. 20,
pp. 570-577, pls. 87-88.

WICKENDEN, R. T. D.
1933. Jurassic Foraminifera from wells in Alberta and Saskatchewan.

Trans. Roy. Soc. Canada, ser. 3, sect. 4, vol. 27, pp. 157-170, pls. 1-2.

EXPLANATION OF PLATES
PLATE I. Jurassic, Neocomian Triplasia

Figs. 1-8. Triplasia kingakensis Loeblich and Tappan, new species......
I, Side view of flattened paratype (U.S.N.M. No. P267a) showing
Flabellammuina-like character. 2a, Side view of quadrate megalo-
spheric paratype (U.S.N.M. No. P267b) showing large planispiral
coil, and the less-prominent fourth angle. 2b, Top view of quadrate
specimen showing rounded aperture and less-prominent fourth
angle. 3, Side view of elongate and narrow paratype (U.S.N.M.
No. P267c). 4a, Side view of holotype (U.S.N.M. No. P266),
showing rapidly flaring test, and small coil of the microspheric
generation. 4b, Top view of holotype showing moderately exca-
vated sides, slightly lower third angle, and rounded aperture. 5,
Side view of smaller paratype (U.S.N.M. No. P267d) showing
large megalospheric coil. 6, Side view of paratype (U.S.N.M.
No. P267e) showing loss of third angle in the final chamber with
reversion to the Flabellammina type of development. 7, Side view
of paratype (U.S.N.M. No. P267f) showing Flabellammina-like
form, with a third angle suggested in the terminal chamber. 8, Side
view of paratype (U.S.N.M. No. P267g) with well-developed coil
and later triangular portion. All & 27. From the Lower Jurassic
Kingak formation, northern Alaska.

Fig. 9. Triplasia bartensteini Loeblich and Tappan, new species..........
Side view of holotype, showing deeply excavated sides and flaring
test (after Bartenstein and Brand), X 30. Middle Jurassic (Dogger
Epsilon) of northwestern Germany.

ic. (iG rilasws elegans CM yatlivke) i. of ieee decree saiacceaealcleies siete amas
Side view of holotype, showing narrow, elongate test, slight exca-
vation of sides, and broadly rounded angles (after Mjatliuk), 30.
Upper Jurassic of Saratov District, U.S.S.R.

Piss: ilasboucesplasia qurassica (Mi jatlitl:) 2% os t.. oisiciec 2 vietwleltiole we calenee
Ita, Side view of holotype, showing flattened character, with the
third angle developed only in the early portion of the test. 11b,
Edge view (after Mjatliuk), & 30. Upper Jurassic of Saratov Dis-
trict) U-SrSsk.

Figs. 12a-16. Triplasia commutata (Loeblich and Tappan)...........+....
12a, Side view of holotype (U.S.N.M. No. 106017) showing large
planispiral coil and more narrow triangular uniserial portion. 12b,

' Top view, showing rounded aperture, X 27, from the Rierdon for-
mation (Upper Jurassic, Callovian) of Wyoming. 13, Side view of
young paratype (U.S.N.M. No. 106021) showing coil and first
triangular chamber, X27, from the Rierdon formation of Mon-
tana. 14, Side view of hypotype (U.S.N.M. No. P885), X 22,
from the Redwater formation (Upper Jurassic, Oxfordian) of
Wyoming. 15, Side view of narrow microspheric paratype

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Page
(U.S.N.M. No. P880), X22, from the Rierdon formation of
Wyoming. 16, Side view of hypotype (U.S.N.M. No. P8o4),
< 22, from the Swift formation (Upper Jurassic, Oxfordian)
of Montana.

Figs. 17a-21b. Triplasia georgsdorfensis (Bartenstein and Brand)........ 12
17a, Side view of “holotype” of JT. emslandensis emslandensis
showing large irregular coil, and triangular later portion. 17b,
View of side showing regularly arched sutures. 17c, Cross sec-
tion (after Bartenstein and Brand), & 25, Upper Valendian of
Germany. 18, Side view of small hypotype of “Tetraplasia
georgsdorfensis” (U.S.N.M. No. P896a) with almost obsolete
coil, quadrate section, and slightly tapering final chamber, X 22,
from the upper Valendian of Germany. 19a, Side view of para-
type of “Triplasia emslandensis emslandensis,’ showing more
elongate form, with final chamber tapered as in the preceding
quadrate form. 19b, Top view (after Bartenstein and Brand),
X< 35, from the lower Hauterivian of Germany. 20a, Side view
of hypotype of “Triplasia emslandensis emslandensis” (U.S.N.M.
P8907) showing the distinct similiarity of the triangular and
quadrate specimens. 20b, Top view, 22, from the lower Hau-
terivian of Germany. 21a, Side view of quadrate hypotype
(U.S.N.M. No. P&96b) showing early planispiral coil and quad-
rate later chambers, gently arched sutures, nearly parallel sides
and distinct similarity of shape of chambers to those of the preced-
ing triangular figured specimen. 21b, Top view, showing some-
what less excavation of the sides than figure in 20, but approxi-
mately the same as in figure 19, X 22, from the upper Valendian
(Lower Cretaceous) of Germany.

Piate 2. Neocomian, Albian Triplasia

Figs. ta-4b. Triplasia pseudoroemeri Bartenstein and Brand.............. 15
1a, Side view of holotype, showing straight sides, high cham-
bers, and rounded angles. 1b, Top view, showing nearly flat sides
(after Bartenstein and Brand), X 30, from the upper Valendian
of Germany. 2a, Side view of quadrate form, the holotype of
“Tetraplasia quadrata,’ showing similarity to the triangular
form in the much-reduced coil, parallel sides, very slightly
arched sutures, broad rounded angles, and nearly flat sides.
2b, Top view, showing ovate aperture (after Bartenstein and
Brand), 25, upper Valendian of Germany. 3a, Side view of
paratype of T. pseudoroemeri. 3b, Cross section of early por-
tion showing nearly flat to slightly convex sides. 3c, Top view,
showing slightly concave sides of the later portion, and ovate
aperture (after Bartenstein and Brand), X 30, upper Valendian-
lower Hauterivian of Germany. 4a, Side view of juvenile form
figured by Eichenberg, showing similarity to the types of this
species. 4b, Top view (after Eichenberg), X 25, Hauterivian of
Germany.

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN

Bissm5a-cComiatplasauorosserugosa ten) Warmtnacrce occ aemcineeomeie oe:
sa, Side view of holotype, showing coarsely arenaceous test. 5b,
Face view showing very slightly arched sutures and elevated
final chamber. 5c, Top view, showing nearly flat sides and ovate
aperture (after ten Dam), 30, Hauterivian of the Netherlands.

BisseOasbweiriplastasacuia battenstein) and Brande. 4. sene eee eee ee
6a, Side view of holotype, showing sharp angles, slightly flaring
test, and moderately arched sutures. 6b, Top view, showing flange-
like angles, excavated sides, and slitlike aperture (after Barten-
stein and Brand), X 30, upper Valendian of Germany.

Figs. 7a-13b. Triplasia glenrosensis Loeblich and Tappan, new species....
7a, Side view of holotype (U.S.N.M. No. P899), a large tri-
angular specimen. 7b, Top view, showing slitlike aperture. 8, Side
view of paratype (U.S.N.M. No. Poooa) showing triangular
early development of the uniserial chambers and loss of the third
angle in the later stages, with reversion to a Flabellammina-type
of growth. 9, Side view of paratype (U.S.N.M. No. Pgoob)
showing planispiral coil, triangular development, and rounded ter-
minal chamber which has lost the angularity, showing a tendency
toward the Ammobaculites type of growth. 10, Side view of
quadrate paratype (U.S.N.M. No. Poooc) showing one side
somewhat narrower than the opposite. 11, Side view of paratype
(U.S.N.M. No. Poood) which has lost the angularity in the final
chamber. 12, Side view of small triangular paratype (U.S.N.M.
No. Poooe). 13a, Side view of quadrate paratype (U.S.N.M.
No. Pooof). 13b, Top view, showing quadrate section and rounded
aperture. All & 22. From the Glen Rose formation (Lower Cre-
taceous, Albian) of Texas.

Figs. 14a-20. Triplasia goodlandensis (Cushman and Alexander)........
14a, Side view of quadrate topotype (U.S.N.M. No. Poo2a). 14b,
Top view, showing moderate excavation of the sides. 15, Side
view of small megalospheric topotype (U.S.N.M. No. Poo2b). 16,
Side view of small, narrow, megalospheric topotype (U.S.N.M. No.
Poo2c). 17, Side view of small megalospheric topotype (U.S.N.M.
No. Pgo2d) showing large coil. 18, Side view of small micro-
spheric topotype (U.S.N.M. No. Poo2e) showing more flaring
test. 19, Side view of larger microspheric topotype (U.S.N.M.
No. Poo2f) showing early coil, flaring test and very strongly
recurved sutures. 20a, Side view of large microspheric topotype
(U.S.N.M. No. Poo2g) showing planispiral coil, with a few
flattened Flabellammina chambers before the triangular develop-
ment and the rapid flaring of the triangular portion. 20b, Top
view, showing acute angles and rounded aperture. All X 48.
From the Goodland formation (Lower Cretaceous, Albian) of
Texas.

Figs. 21-22b. Triplasia mexicana Loeblich and Tappan, new species......
21, Side view of small paratype (U.S.N.M. No. P1023) showing
small coil and gently arched sutures. 22a, Side view of holotype
(U.S.N.M. No. P1022) showing flared test. 22b, Top view,

Page
14

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Page
showing moderately excavated sides and ovate aperture. 22%
From the Barrill Viejo shale (Lower Cretaceous, Hauterivian)
of Coahuila, Mexico.
PiaTE 3. Albian Triplasia
Figs. 1-13. Triplasia incerta (Alexander and Smith).................0.- 21

I, Side view of hypotype (U.S.N.M. No. Po931), showing typi-
cal Flabellammina-like early portion, with third angle developed
in the last one-third of the test, X10. From the Fort Worth
formation (Lower Cretaceous, Albian) of Texas. 2, Side view
of hypotype (U.S.N.M. No. Po25a) from the Duck Creek forma-
tion (Lower Cretaceous, Albian) of Oklahoma, showing elongate
Flabellammina stage, X22. 3, Side view of broad and elongate
hypotype (U.S.N.M. No. Po930) from the Kiamichi formation
(Lower Cretaceous, Albian) of Texas, with very short triangular
portion, X 22. 4, Side view of hypotype (U.S.N.M. No. P933)
showing an unusually long triangular portion and narrow test, from
the Fort Worth formation of Texas, 22. 5, Side view of hypotype
(U.S.N.M. No. Po932) from the Fort Worth formation of Okla-
homa, X10. 6, Side view of hypotype (U.S.N.M. No. Po28a)
from the Denton formation (Lower Cretaceous, Albian) of
Texas, X 22. 7a, Side view of holotype (U.S.N.M. No. Pg2q4)
from the Fort Worth formation of Texas. 7b, Edge view, show-
ing compressed test and low ridgelike third angle. 7c, View of
rounded aperture. X22. 8a, Side view of quadrate hypotype
(U.S.N.M. No. Po926) from the Weno formation (Lower Cre-
taceous, Albian) of Texas, showing coil and early development of
third angle. 8b, Edge view, showing insertion of fourth angle ap-
proximately halfway up the test. 8c, Top view, showing elongate
aperture. XX 22. 9, Side view of elongate hypotype (U.S.N.M.
No. P929) from the Kiamichi formation of Texas, < 22. 10, Side
view of small hypotype (U.S.N.M. No. P927) from the Fort
Worth formation of Texas, X 22. 11, Side view of small hypo-
type (U.S.N.M. No. Po25b) from the Duck Creek formation
of Oklahoma showing more rapid development of third angle,
X 22. 12, 13, Side views of hypotypes (U.S.N.M. Nos. Po28b, c)
from the Denton formation of Texas, x 22.

Figs. 14-20c. Triplasia rugosa (Alexander and Smith).................5- 27
14, Side view of hypotype (U.S.N.M. No. Pgo55) showing well-
developed planispiral coil, later triangular uniserial portion with
final chamber tending to become rounded. 15a, Side view of tri-
angular hypotype (U.S.N.M. No. Po56a). 15b, Top view. tI6a,
Side view of quadrate hypotype (U.S.N.M. No. Po56b). 16b,
Top view. Figures 14-16 X 22. From the Duck Creek formation
of Oklahoma. 17a, Side view of compressed topotype (U.S.N.M.
No. Po954a), showing Flabellammina-like character also found in
the holotype. 17b, Top view. 18a, Side view of topotype (U.S.N.M.
No. Po54b) showing triangular development of the final chamber.

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 57

Page

18b, Edge view. 19, Side view of topotype (U.S.N.M. No. Po54c)
with well-defined planispiral coil, followed by a flattened uniserial
portion which becomes triangular in the last two chambers. Figures
17-19 X27. 20a, Side view of holotype (U.S.N.M. No. Po52)
showing flattened development which led to its original descrip-
tion as a Flabellammina. 20b, Edge view showing degree of com-
pression. 20c, Top view, showing slightly elongate aperture. 22.
Figures 17-20 from the Duck Creek formation of Texas.

Pirate 4. Albian Triplasia

Figs. 1-9. Triplasia acutocarinata (Alexander and Smith)............... 19
1, Side view of hypotype (U.S.N.M. No. Pg906) from the Kiamichi
formation of Texas. 2, Side view of unusually broad hypotype
(U.S.N.M. No. P1025) from the Fort Worth formation of Okla-
homa. 3, Side view of large flaring hypotype (U.S.N.M. No. Pgo7)
from the Duck Creek formation of Texas. 4, 7, Side view of hypo-
types (U.S.N.M. Nos. Pooga,b), from the Weno formation of Texas.
5a, Side view of topotype (U.S.N.M. No. P904), showing early
planispiral coil and produced final chamber. 5b, Top view, showing
a more elongate aperture than is typical. 6a, Side view of holo-
type (U.S.N.M. No. Po03). 6b, Top view, showing flangelike
angles and rounded aperture. Figures 5 and 6 from the Duck
Creek formation of Texas. 8a, Side view of quadrate hypotype
(U.S.N.M. No. Poogc). 8b, Top view, showing the narrow angles
typical of the species, and the rounded aperture. From the Weno
formation of Texas. 9, Side view of long and narrow hypotype
(U.S.N.M. No. Po908) from the Denton formation of Texas. All
figures XX 22.

Figs. 10-15b. Triplasia wenoensis (Alexander and Smith).............. 28
10-12, Side views of hypotypes (U.S.N.M. Nos. Po959 a-c), show-
ing small size, flaring test, and roughly finished wall. 13, Side view
of larger hypotype (U.S.N.M. No. Po58). 14a, Side view of holo-
type (U.S.N.M. No. P957), showing small size, rounded angles,
and moderately excavated sides. 14b, Top view, showing rounded
aperture. 15a, Side view of quadrate hypotype (U.S.N.M. No.
Pg50d). 15b, Top view, showing moderately excavated sides. All
figures X 22. From the Weno formation of Texas.

Pirate 5. Albian to Senonian Triplasia

Figs. 1a-7. Triplasia insolita (Alexander and Smith).................06. 24
Ia, Side view of holotype (U.S.N.M. No. Po934), showing tri-
angular character of early portion grading into a quadrate later por-
tion. 1b, Top view, showing rounded aperture. 2a, Side view of
topotype (U.S.N.M. No. P936a), showing early planispiral coil and
later Flabellammina-stage before development of the third angle.
2b, Edge view, showing flattened character of coil and early arched
chambers with abrupt thickening in the later development. 3a, Side
view of topotype (U.S.N.M. No. Po936b), showing quadrate de-

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Page
velopment immediately after the planispiral stage. 3b, Edge view,
showing compressed early portion and abrupt thickening. Figures
1-3 from the Duck Creek formation (Lower Cretaceous, Albian) of
Texas. 4, Side view of small hypotype (U.S.N.M. No. Po4o) from
the Weno formation (Lower Cretaceous, Albian) of Texas. 5, Side
view of hypotype (U.S.N.M. No. Po938a) from the Main Street
formation (Lower Cretaceous, Albian) of Texas. 6, Side view of
hypotype (U.S.N.M. No. Po39) from the Fort Worth formation
(Lower Cretaceous, Albian) of Texas. 7, Side view of hypotype
(U.S.N.M. No. P938b) from the Main Street formation of Texas.

All figures X 22.

Figs. 8-15. Triplasia nodosa Loeblich and Tappan, new species........... 25
8, Small triangular paratype (U.S.N.M. No. Po44a) showing early
coil and produced final chamber, from the Grayson formation
(mid-Cretaceous, Cenomanian) of Texas. 9, 10, Larger triangular
paratypes (U.S.N.M. Nos. Po42a-b) from the Grayson formation
of Oklahoma, with the latter specimen losing the third angle in the
later chambers. 11, Paratype (U.S.N.M. No. Po49), showing early
coil followed by a flattened uniserial portion, the triangular develop-
ment appearing only in the last half of the test. From the Main
Street formation (Lower Cretaceous, Albian) of Texas. 12a, Side
view of holotype (U.S.N.M. No. Po41), showing typical triangular
development and extended neck. 12b, Top view, showing triangular
section. From the Grayson formation of Oklahoma. 13a, Quad-
rate paratype (U.S.N.M. No. P947) showing constricted sutures
and nodose appearance of the pyriform later chambers. 13b, Top
view, showing quadrate section. From the Main Street formation
of Texas. 14, 15, Triangular paratypes (U.S.N.M. Nos. Po44b-c),
showing angular nodose tests and much-produced neck. From the
Grayson formation of Texas. All figures X 22.

Ie = MMe TiAl OOoOiee INSERSS declggobdssonododebuodaceéanse- 30
16, Face view of narrow paratype, showing gently arched sutures.
17a, Side view of holotype, a flared specimen. 17b, Face view. 17¢,

Top view, showing slightly excavated sides, subacute angles, and
rounded aperture (after Reuss). Approximately 10. Lower
Senonian or Turonian of Austria.

LENESISS itso, Wi awelomten tocol. (MMS) oo Sbodochoscuseadcdsoocenacocbacc 32

18, 19a, Side views of typical triangular hypotypes (topotypes of
Haplophragmium murchisoni of Beissel (U.S.N.M. Nos. P0963,
P964), showing regular planispiral coil and gradually enlarging
test. 19b, Top view, showing slight excavation of sides, rounded
angles, and slit aperture. Both X10. From the Mucronaten-
Kreide (upper Senonian) of Germany. 20a, Side view of quadrate
hypotype (U.S.N.M. No. P0967), showing basal coil, very slightly
enlarging test, and gently arched sutures. 20b, Top view, showing
quadrate section, excavated sides, and slit aperture. X22. From
the lower Senonian of Germany.

Figs; 2ta-22b,Triplasiawomers Reuss) k see ee eee ae ee ee 38
21a, Side view of holotype, showing rounded base, nearly parallel

NO. I5 TRIPLASIA REUSS—LOEBLICH AND TAPPAN 59

Page

margins and sharp angles. 21b, Face view, showing weakly curved
sutures, and comparatively high chambers. 21c, Top view, showing
somewhat convex sides of the later portion and the rounded aper-
ture (after Reuss), approximately 16. Mucronaten-Kreide
(upper Senonian) of Germany. 22a, Side view of hypotype
(U.S.N.M. No. Po988), showing basal coil, slightly curved test
with nearly parallel margins, and high narrow chambers. 22b, Top
view. X22. From the upper Senonian of Germany.

Puiate 6. Senonian Triplasia

Figs. 1-5. Triplasia deadericki Loeblich and Tappan, new species........ 35
1, Side view of compressed paratype (U.S.N.M. No. Po81), which
is Flabellammina in character, showing low early chambers and the
higher and rounded final chamber. 2, Side view of paratype
(U.S.N.M. No. Po79a) with planispiral early portion, followed by
triangular uniserial portion, and last chambers becoming higher and
rounded in section. 3a, Side view of holotype (U.S.N.M. No. Po78),
showing typical triangular form with moderately excavated sides and
higher rounded final chamber. 3b, Top view, showing ovate aper-
ture, and excavated sides of triangular portion. 4a, Side view of
small quadrate paratype (U.S.N.M. No. Po7zob), showing early
coil and comparatively low early quadrate chambers. 4b, Top view,
showing quadrate section. 5, Side view of small paratype
(U.S.N.M. No. Pozoc) with rounded final chamber. All x 22.
From the Annona chalk of Arkansas.

Figs. 6-13. Triplasia taylorensis (Cushman and Waters)..............06. 30
6, Side view of hypotype (U.S.N.M. No. Pioooa) showing a typi-
cal large flaring specimen, with broad low chambers. 7a, Side view
of hypotype (U.S.N.M. No. Prooob) which is coiled at the base,
then triangular in section, and becomes quadrangular in the last half
of the test. 7b, Top view, showing quadrate section. Figures 6-7
10. From the upper Taylor marl of Texas. 8, Holotype (Cush-
man coll. No. 12033), a smaller form, which is otherwise typical of
the species, X 22. From the Taylor marl of Texas. 9a, Side view
of small quadrate hypotype (U.S.N.M. No. 1o00c), showing typical
flaring test. 9b, Top view showing sharp angles, excavated sides,
and slit aperture. 10a, 11, 12, Side view of regularly flaring triangu-
lar hypotypes (U.S.N.M. Nos. Pioood-f). 1ob, Top view, show-
ing excavated sides and slit aperture. Figures 9-12, X I0. 13, Side
view of small quadrate hypotype (U.S.N.M. No. Prooog), X 22.
Figures 9-13 from the upper Taylor marl of Texas.

Figs. 14-19b. Triplasia plummerae Loeblich and Tappan, new species...... 36
14, Side view of paratype (U.S.N.M. No. Po84a), showing flaring
triangular test, broad and low early chambers, and higher and some-
what rounded final chamber. 15a, Side view of quadrate paratype
(U.S.N.M. No. Po84b), showing gently arched sutures. 15b, Top
view, showing flat to moderately excavated sides. 16, Side view of
triangular paratype (U.S.N.M. No. Po84c). 17a, Side view of quad-

Figs. 1a-5. Triplasia cushmani (Alexander and Smith)

Figs. 6a-9. Triplasia rugosissima (Alexander and Smith)

Figs. 1oa-11b. Triplasia saratogensis Loeblich and Tappan, new species...

Figs. 12a, b. Triplasia species

Figs. 13a-14b. Triplasia fundibularis (Harris and Jobe)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. [17

rate paratype (U.S.N.M. No. Po84d). 17b, Top view, showing
nearly flat sides and slit aperture. 18, Side view of paratype
(U.S.N.M. No. Po84e). 19a, Side view of holotype (U.S.N.M.
No. P9083), showing broad and low early chambers, and higher,
rounded final chamber with slight neck. 19b, Top view, showing
nearly flat sides and slit aperture. All X 10. From the upper Tay-
lor marl of Texas.

Pate 7. Senonian to Pliocene Triplasia

1a, Hypotype (U.S.N.M. No. Po969) showing early planispiral coil
and triangular later portion. 1b, Top view, showing acute angles and
excavated sides and slit aperture. From the Annona chalk (Upper
Cretaceous, Senonian) of Arkansas. 2, Side view of coarsely
arenaceous holotype (U.S.N.M. No. Po89), showing early coil and
broad low triangular chambers, from the Pecan Gap (Upper Cre-
taceous, Senonian) of Texas. 3a, Side view of large quadrate hypo-
type (U.S.N.M. No. Poor), 3b, Top view, showing excavated
sides and slit aperture. From the Annona chalk of Arkansas. 4, 5,
Small hypotypes (U.S.N.M. Nos. Po71a-b) from the Ozan sand
(Upper Cretaceous, Senonian) of Arkansas. All X 22.

6a, Side view of quadrate hypotype (U.S.N.M. No. Pgg7a). 6b,
Top view, showing nearly flat sides and slit aperture. From the An-
nona chalk of Arkansas. 7a, Side view of holotype (U.S.N.M.
No. P9093), showing coarsely arenaceous test, triangular uniserial
portion, and somewhat flattened final chamber. 7b, Top view,
showing slit aperture. From the Taylor clay of Texas. 8, 9, Side
view of hypotypes (U.S.N.M. Nos. Poo7b-c) from the Annona
chalk of Arkansas. All > Io.

toa, Holotype (U.S.N.M. No. Pioo5), showing typical triangular
form, and subparallel margins. tob, Top view, showing slightly
convex sides and broadly rounded angles, and ovate aperture. IIa,
Side view of quadrate paratype (U.S.N.M. No. P1006), showing
moderately excavated sides in the early portion and nearly flat sides
in the later portion. 11b, Top view, showing flattened sides, quad-
rate section, and ovate aperture. From the Saratoga chalk (Upper
Cretaceous, Maestrichtian) of Arkansas. X 22.

12a, Side view of specimen (U.S.N.M. No. 560500), showing early
coil and later triangular portion. 12b, Top view, showing slit aper-
ture. X 22. From the Lodo formation (Paleocene and Eocene) of
California.

13a, Side view of large hypotype (U.S.N.M No. Prorga), showing
broad, low chambers, flaring test, and excavated sides. 13b, Top
view. 14a, Side view of smaller hypotype (U.S.N.M. No. Pior4b),

)

Page

NO. 15 TRIPLASIA REUSS—LOEBLICH AND TAPPAN

showing broadly rounded base and gently arched sutures. 14b, Top
view. X27. Topotypes from the Midway formation (Paleocene)
of Arkansas.

Heeets) rina aurmasicus (Weller) 6 .a64.d2 es gag hou ae veel amorous
Holotype, showing early planispiral coil and triangular uniserial
portion, with rounded angles (after Keller). Approximately x 27.
From the lower Senonian (Campanian) of the western Caucasus,
RES Sate

Ex SammOa 17 Dum h7ay Ste IMUIUECGINEUISS)) = aici sie) «rs,<rciasysi se crelelereicioeiectoeieee
16a, Face view of holotype, showing absence of a distinct coil and
gently arched uniserial chambers. 16b, Side view, showing grooved
angles described by Reuss, but which may be due to breakage of the
fragile, very acute angles. 16c, Top view, showing very sharply
angled test and deeply excavated sides. 17a, Face view of more
flared paratype. 17b, Side view (after Reuss). From the Miocene
of Poland.

Figs. 18a-b. Triplasia wrightt (De Amicis)...........ccdeeccccesccvdes
18a, Side view of holotype, showing large coil and comparatively
narrow triangular uniserial portion with nearly parallel margins
and broad, low, slightly arched chambers. 18b, Top view, showing
slightly convex sides and ovate aperture (after De Amicis), x 16.
From the lower Pliocene of Italy.

Prate 8. Oligocene to Recent Triplasia

Higsnaarc.\ Leipiasie MUnGOrice ( MAaIZOM) Setas.< cd gsc ia:s.c sie wiareieres.nie seg auncage
1a, Side view of quadrate holotype, showing large planispiral coil
and broad, low, quadrate uniserial chambers. 1b, Face view, show-
ing compression of the planispiral coil and arched sutures of the
uniserial portion. 1c, Top view, showing excavated sides and ovate
aperture (after Majzon). X 28. From the Oligocene of Hungary.

Figs. 2-4b. Triplasia marwicki Loeblich and Tappan, new species......
2, Side view of small paratype (U.S.N.M. No. 1018a), showing
flared test and arched sutures. 3a, Side view of larger paratype
(U.S.N.M. No. Pio18b). 3b, Top view. Both from the Miocene
of Motatura, New Zealand. 4a, Side view of holotype (U.S.N.M.
No. Prio16). 4b, Top view. From the Miocene of Kawhia, New
Zealand. All X15.

Bagse5-opy feripiosia: vartauies (Brady) 2.) a0 ..<siiclees se aalslaweieie als eaweete

5, Side view of small quadrate hypotype (U.S.N.M. No. Pio29a).
6, Side view of small triangular hypotype (U.S.N.M. No. Pio29b).
7a, Side view of large triangular hypotype (U.S.N.M. No. Pro29c).
7b, Top view showing sharp angles, concave sides, and slit aperture.
8, Side view of small hypotype (U.S.N.M. No. Pio29d), 9a, Side
view of large quadrate hypotype (U.S.N.M. No. Pro29e), showing
arched sutures. ob, Top view, showing quadrate section and slit
aperture. All 15. From coral sands at 210 fathoms, off Kandavu,
Fiji Islands.

Page

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 17 INOAIS.PE at

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 117, NO! 15, PES2

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

117, NO. 15, PL. 3

VOL.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 15, PL. 4

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOESII7; sNOMIipaPie5

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 15, PL. 6

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

| iMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 15, PL. 7

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS

FORAMINIFERAL GENUS TRIPLASIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 16

Roebling Fund

SOLAR VARIATION AND PRECIPITATION
AT PEORIA, ILLINOIS

BY
Cc. G: ABBOT

Research Associate, Smithsonian Institution

(Pustication 4095)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 3, 1952

The Lord Baltimore Preos

BALTIMORE, MD., U. 8 A.

oye
eca? ¥

Roebling Fund

SOLAR VARIATION AND PRECIPITATION AT
PEORIA, ILLINOIS

By C. G. ABBOT
Research Associate, Smithsonian Institution

In a recent paper * it was shown that the intensity of the sun’s radia-
tion, as it is outside the earth’s atmosphere, varies simultaneously in
23 regular periods, all aliquot parts of approximately 272 months.
It would naturally follow that details of variation in observed values
of the solar constant of radiation would tend to repeat at intervals of
about 23 years. Figure 4A of the paper cited shows that this is indeed
the case.

Many years ago it was shown that the precipitation at Peoria, IIl.,
also tended very strongly to exhibit repetitions of features at intervals
of 23 years. I reproduce here as figure 1, the figure 33 of an earlier
paper.’

Intending to trace the effect of the solar variations above noted on
weather, it seemed well to study first the records of precipitation at
Peoria, as these were already known to exhibit influences of the
master cycle of about 23 years. I have been engaged over 3 years in
this investigation, for unexpected complexities kept cropping up,
which made it necessary again and again to scrap all results and begin
at the beginning. Without being tedious, it may be said that I have
tabulated over 1,000 months of precipitation records, of the years
between 1856 and 1939, separately in 20 to 30 supposed periods, and
repeated the whole tabulation no less than 14 times.

Futility of determining periods from weather records—Knowing
that the sun’s radiation varies in regular periods, which are integral
submultiples of about 23 years, but being restricted by available solar-
constant observations to little more than 20 years with many gaps and
inaccuracies in these records, I hoped at first to use the continuous
series of Peoria precipitation records from 1856 to 1939 to establish
which submultiples of 23 years are active, and to fix their exact
lengths.

1 Smithsonian Misc. Coll., vol. 117, No. 10, 1952.
2 Smithsonian Misc. Coll., vol. 94, No. 10, 1935.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 16

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOI, 7

It was soon disclosed that the phases of periodicities, though in-
variable in solar variation, shifted in Peoria precipitation records,
depending on the time of the year. I suppose the cause to be associated
with considerations of lag in response to solar impulses. It is patent
that though the sun’s heating potential reaches a maximum daily at
noon, maximum temperatures do not come until several hours after
noon, and their arrival displays different lags in different localities, and
in the same locality at different times and seasons. Similar it must
be with regard to all weather responses to all solar periodicities.

Subdivision of the year—Accordingly, one of my first modifica-
tions of procedure was to prepare separate tabulations for each perio-
dicity in three fractions of the year, namely: January to April, May
to August, September to December. Doubtless a finer subdivision of
the year would theoretically be better, but for periodicities of Io to 20
months in length the number of columns in a tabulation becomes few.

Interference between periodic effects —It is plain that the determi-
nation of every periodic fluctuation in precipitation must be affected by
the presence, implicitly in the data, of above 20 other periods of dif-
ferent lengths. Theoretically these disturbing influences can only be
completely sifted out of the result for one period, if the mean values
are computed from an infinite number of repetitions of the chosen
period. When, instead of infinity, one becomes limited to a few repe-
titions, the computed march of the investigated periodicity must evi-
dently be very imperfectly determined. Hence, on account of paucity
of data, it was impracticable to divide the years beyond the triple
division mentioned above.

Sunspot frequency affects phases.—But still another limitation of
freedom in tabulation soon appeared. The phases of features in the
periodicities computed were found different in different years. This
was found to be associated with sunspot frequency. As said above,
the cause of the phenomena of phase changes is probably to be as-
signed to variable lag of response in weather to solar impulses, depend-
ing on the condition of the atmosphere. Such changes in lag are evi-
dently attributable to the varying character of the atmosphere at dif-
ferent seasons of the year as regards transparency, cloudiness, and
other conditions. But sunspots continually bombard the atmosphere
with multitudes of electric ions. These act as centers of condensation
for the constituents of haze. Hence it was only to be expected that
phase changes would be associated with Wolf sunspot numbers. Here,
again, a compromise had to be made. It was decided to subdivide the
precipitation data in two groups, for Wolf numbers = 20.

PRECIPITATION AT PEORIA—ABBOT

NO. 16

‘uoryeydioe1d el103g ur Ayorporsiod I1eaA-3914}-AJUIMI—I “DIJ

Min
l

AMS

arr

Be Pan
ir
Pale aes

Sey
Zia

TWWYON LN39d3d

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL; Diz,

Necessary roughness of determinations——Between the accidental
influences associated with rainfall, cloudiness, haziness, etc., the un-
determinable interferences of some 20 periodic solar variations on the
periodicity being considered, and the necessary subdivision of the data
for seasonal and sunspot alterations of atmospheric quality, the de-
terminations of periodicities in precipitation became too rough to
warrant conclusions that such and such submultiples of 23 years were
real, effective periods in solar variations.

Length and number of periods necessarily determined from solar
observations.—Thrust back on the solar-constant observations as the
only sound source of knowledge of the solar periods, I laid aside the
investigation of Peoria precipitation to prepare the paper above cited
(footnote 1). In that paper are listed 23 periods that appear to be
real in solar variation, and 15 others that were tried and found want-
ing. In the latter part of the Peoria tabulations the investigation was
limited to the confirmed 23 periods. However, for convenience in
tabulating, the lengths of some periods were slightly altered.

Faulty normals.—Still another unsuspected hitch occurred in the
investigation, which caused several weeks delay and a completely
new tabulation of the precipitation data. This is explained in another
recent paper.® In substance it amounts to this: The monthly normal
values printed at the bottom of the pages in World Weather Records #
make no distinction between times of sunspot maximum and times of
sunspot minimum. Two consequences result. First, the average of the
monthly data computed as percentages of the published normals of
precipitation at Peoria is about 9 percent higher for times when Wolf
sunspot numbers exceed 20 than for times when these Wolf numbers
are below 20. Second, far more serious, and indeed fatal to success
in my investigations made theretofore, is the fact that when averages
of data timed for sunspots 2 20 are separated, there is found a large
discrepancy, month by month, between these averages for Wolf
numbers <20 and for Wolf numbers >20. This plays havoc with
tabulations of periodicity. The published normals, in fact, combine
two contradicting sets of data as foreign to each other in attitude as
dogs are from cats, and tabulations including both sorts in the same
periodicity are worthless.

Final preparation of the data.—The recorded monthly values of
precipitation at Peoria for the years 1856 to 1950, found in World
Weather Records and later publications, were segregated in two

8 Smithsonian Misc. Coll., vol. 117, No. 11, 1952.
4 Smithsonian Misc. Coll., vols. 79, 90, and 105.

NO. 16 PRECIPITATION AT PEORIA—-ABBOT 5

groups, for Wolf numbers 2 20. Monthly normal values were com-
puted separately, 52 years in the high-sunspot group, 30 years in the
other. Two years were omitted, when the months were about evenly
divided between the two classes.

The new normals for Peoria precipitation, as expressed in inches,
are as follows:

Sunspots Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
> 20 1-04) 1.07) 2.50) »3:02 13-00 4-04. 3:70) 3:06 3°56) (2:22) 2946) Or
<20 TGoi tant, pr 2o2 $2.70" B88) iet24) | (3.40!) 20661) ZI8s) “ako a2 zeu eee

From these new normals percentages of normal precipitation were
computed for every month, 1856 to 1950. There are such large and
suddenly changing jumps from month to month in these percentages
of normals that the data were then smoothed by 5-month consecutive
means. Thus, for March compute

Jan. + Feb. + Mar.+Apr.+ May
5

For April drop January and add June, and so on. Even then, as some
exceptionally rainy months occurred, the smoothed data would soar
above 200 percent for a brief interval. With the necessarily few
columns in a tabulation, such an exceptional case might quite vitiate
the determination of a periodicity. Hence for some 20 exceptional
months out of over 1,000, the original values were scaled down to
200 before the smoothing process was done.

Division of the interval—While the number and lengths of the
periodicities to be sought in Peoria precipitation had been fixed by
study of solar-constant observations, and nothing was to be learned
as to the reality or the exact length of these periodicities in solar radia-
tion from the precipitation data, it still was desirable to tabulate
Peoria precipitation in several parts independently. Certain of the
solar periodicities might produce such weak responses in precipitation
that the several groups might give no agreeing results distinguishable
from accidental error. Such weak periodic effects could be neglected.
Such indeed proved to be the case with periods of 27 and 3,'; months,
and with the periodicity of 68 months. There were strong features
in the separate determinations of the periodicity of 68 months, but
they were so confused by overriding interference from various shorter
periods as to be uncertain. Hence that period was dropped.

More important, however, just as the changing atmospheric con-
ditions were found to alter phases with the seasons, and with the
prevalence of sunspots, it proved that, apart from these already noted

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

phase alterations, there were secular changes of phases. One group
of years, though yielding the same forms of periodicities as another,
yielded them in different phases. Such effects were found to vary in
character for periods of different lengths.

In the tabulations, therefore, the years 1856 to 1900 were tabulated
separately from the years 1900 to 1939. Discordant phases appearing
in the earlier interval were shifted to accord with the phases for the
years 1900 to 1939. Then the two determinations were averaged to-
gether. So the full strength of the data from 1856 to 1939 was con-
centrated in the phases prevailing from 1900 to 1939.

Fractional months in periods.—In order to preserve the exact period
in such cases as 92, 114, and 15% months, and others not exactly even
months in length, some columns in tabulating were made a month
longer or a month shorter than others. In case of long periods,
months were interpolated or cut out of the columns from place to
place as required to make the average length of the columns equal to
the period. Such additions to columns may be noted in tables 1 and 2
for the 93-month periodicity, and in tables 3 and 4 for 15 months.

Examples of tabulation.—In order to fix ideas, I now give two
actual tabulations and corresponding plots, to bring out the process
employed, and to clarify the explanations above. I select the periods
of 9 and 154 months. Table 1 gives the tabulation for 9% months
suited to Wolf numbers >20, and table 2 that for Wolf numbers <20.
In each table the segregation of the year by 4-month intervals is pre-
served. Also, two separate tables for each 4-month interval are made,
one for the first half, the other for the last half of the 84-year in-
terval. As stated above, there is often found a discrepancy in phase
between results of the first and second halves of the 84-year interval.
In table 1, sections A, and B,, the symbol A,4, means that to bring
the phase of the first half, Ai, to that of the second half, B,, the mean
values for A, are moved one month backward. The symbol A,%, ob-
tains in sections A, and B., and A3y3 was required for sections
A; and B;. Similar symbols for shifts appear in the tabulation of the
data suited to Wolf numbers <20. The final columns, marked A,
give the departures from the average percentage of results in the
general means. The use of these columns of departures will be
explained below.

Tables 3 and 4, relating to the periodicity of 15 months, will be
understood from the preceding description of tables 1 and 2 for 9%
months. As before, symbols involving # mean that the means of
the first half of the data were shifted backward by one or more
months, and symbols involving |) indicate the contrary.

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NO. 16

Apr. Jan.
AGO) 9273
127 89
139 = 98
149 112
133 89
116 98
106 97
94 91
94 87
94 84
III 104
87 129
68 124
49 III
49 115
39 «85
Ag

June July
84 04
109 86
127. 80
140 84
139 95
153 93
154 72
I31 67
106 58
Toy aL
87 45
79 «682
90 89
QI 101
96 98
1o8 125
As

Dec. Dec.
262) 367,
137. 55
142 85
135 85
107, 123
Oyo e
Oy WLEL
66 gl
72 98
92 71
96 102
HI3n ig
114 106
104 100
85 108
105 90
102

PRECIPITATION AT PEORIA—ABBOT

TABLE 3.—Peoria precipitation. The 151-month period

Sunspots > 20

Jan. Mean Apr. Mar. Jan.
"92 193) 1070) 106
110. 106.0 120 84 95
100 106.2 103 64 85
II5 114.7 84 89 86
I3h 104.3 66) Sra 60
129 97.0 49 114 63
1160) 94..5 58 106 86
104 93.0 63° nor 198
Z5 9402 89 92 106
69 92.0 87 58 109
70 97.0 103 83. 121
78 105.0 9 98 102
96 101.2 96 106 84
107 89.0 $9 220 73
11g 94.7 79 142 83
124 89.5 73 G22) 877
Mean Aug. June May Aug
704s) 7080) 2277) 228
91.4 Sy ¥22-" ysis’ 13x
104.2 70) 104, 9132) "32
110.4 80 7 4ua7 aah
112.8 66,907) eng) 250
116.0 44 54) 35 tes
106.6 Sy eeess Eos, aS)
93-4 78) 107 "£40! 1130
86.4 87.74 134 29
92.8 100 93 Yx20° a2
87.0 108 103 89 137
92.0 102 104 76 124
95.4 90 109 62 104
99.0 92 101 90 113
104.6 88 87 95 106
110.6 Sr oo) 12 86
97
Nov. Sept. Mean Oct. Nov.
72) 490 795 705
151 76 102.0 27D
162 77 LOAs7 126 101
143 77 101.0 119 94
121 86 = 98.9 121 88
116 78 = 994.4. 110 86
139 82 101.3 96 = 84
120 88 102.1 93 78
11006 86s: 103.7 III 70
I05 8r 105.1 134 82
106 99 115.3 136 77
88 90 116.1 109 «682
QI 86 114.4 116 85
92 105 109.4 83 114
nae LES Gls. 91 88
98 91 109.7 77. «9S
99 90

9
B,
Apr. Feb. Apr. Mean As Mean A
71 726) a6
930 84. 50 87.7. 105.0 96.3. — 2.0
100) = 8&3 30 79.0 YToO1.2, 00.5 — 6.2
106 96 41 83.7 89.0 86.3 —12.0
98 103 75 85.8 94.7 89.7. — 8.6
106 126 97 92.5 89.5 91.0 — 7.3
70 4155) vod » 96.5 ¥06.0° moxr.2 230
68 167 132 104.8 106.2 105.5 + 7.2
62 i78t 148 Ose. Ora7 orr13.6 --153s
63 158 132 I01.2 104.3 102.7 + 4.4
50 136 109 100.3 97.0 98.6 + 0.3
69 124 123 102.0 904.5 98.3 0.0
an 133) TOG) 100.9 93-0 96.7. — 1.6
$3) 120 99 99.0 94.2 96.6 — 1.7
84 149 85 103.7 92.0 97.8 — 0.5
116 «158 88 105.7 97.0 101.3 + 3.0
Mean _— 08.3
Bo
July July Mean AN; Mean A
39 71S
yor IOS er rte7 92.0 t101.8 — 0.2
PS, 317 206.5 95.4 100.9 — 1.1
GA 120°) 101.2 99.0 100. — I.9
73> 13t 0203.7 (104.6 “roger -p ia
Taz 93 94.3 110.6 102.4 + 0.4
110 109 100.3 91.4 95.8 — 6.2
127 114 109.3 104.2 106.7 + 4.7
037 100 10:2 rTlo.45 110.3) --'8.3
145 90 110.0 112.8 107.8 -+ 5.8
139 109 109.5 116.6 113.0 -+11.0
I50 101 109.5 106.6 108.0 -+ 6.0
138 100 100.5 03-4 96.9 — 5.1
129 106 105.2 86.4 95.7 — 6.3
Toy “DLT 99.0 92.8 95.9 — 6.1
87 98 95-5 87.0 91.3. —I0.7
Mean 102.0
Bg
Oct. Oct. Nov. Dec. Oct. Mean Acts Mean
“TA, 7°19 "24 734 538
85 08 65 “1237 68) ‘93:57 stone 97.6
88 78 67 132 85 95-3 102.1 98.7
88 61 67 705 104) Ot. 103.7 97.4
76 = s« tr 72: ¥30) 120 98.3 105.9 102.1
83 89 54 142 135 99.9 115.3 107.6
103. 102 64 126 143 102.6 116.1 109.3
93 102 65 109 133 04.7 114.4 104.5
102 108 82 112 128 101.9 109.4 105.6
128 81 101 86 x12 10556 xTr5.7 10.6
146 aa DLO 88 97 102.4 109.7 106.0
Ler 59 123 88 108 97.I 102.0 99.6
120 59 112 I0or1 90 97.6 104.7 I01.0
I00 70 99 100 71 91.0 I0I1.0 96.0
108 75 96 98 64 886 98.9 93.7
84 68) 92 %73)) 601) 79:7) ota 67.0
67 ean I01.1

a Be a oc aoa
PIN OHROMNW DAHWNW

col

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Graphical presentation—tThe results on the 9$-month periodicity,
in tables 1 and 2, are shown graphically in figures 2 and 3. In each
of the six charts shown in each figure, the lower curve (a), in light

TABLE 4.—Peoria precipitation. The 15$-month period

Sunspots < 20

Ai By
Apr. Feb. Mar. Jan. Apr. Mean Feb. Mar. Jan. Mar. Apr. Mean Ay | s Mean A
Ural AON Sor henO7 me ROS WO) oaks e2ty 43 hie OS
62) arr) Lirrs) ar08') 40) S20 722 98 Iol 86 77 LAO ‘TOTO 81.8 91.4 —10.4
54 O7 S102) sstose errs 94.2 89 108 121 85 113 106.8 89.4 98.1 — 3.7
83° FI5) a2r aos 98 105.0 TiOn 24) o22 Ero 08 If.0 r0g2 “107-5 -- 75.3
76 104 II10 79 «108 95.4 143/122 108 air "108 x«ireate (104;4 108.3) 1-655
79 f23° 126 2 103 4104.6 144 95 I22) X20) (103) “1oO570) “T1064 ) 9107277 ed
88 ror 109 65 100 92.6 i170. 103 216) 2S) W100 ) To9.2) 107-2. | WroSsee ora.
SO) LOO) aat22 54 107 04.4 188 106 Q6) 45) S107) ars 94.2 103.8 -+ 2.0
47 93) JULIO 48 92 78.0 176 88 108 134 92 105.5 105.0 105.2 + 3.4
65 too)" 114: 67 84 86. Tsau) 103) fz) fA 7: 84 I15.0 95-4 105.2 -+ 3.4
70 80 99 54 82 77.0 122) (rzo “239) *138 82 «19.8 104.6 112.2 -+10.4
65 86 92 93 73 81.8 LOLS TIS Wrst 3m 72) Tt2s0 92.6 102.3 + 0.5
80 68 SFA ars 94 89.4 Q4. 24 | 120 7 “104 94 + I10.5 94.4 102.4 + 0.6
905 81 O2) e057) TOL wrogs2 Qo) 122 142 99° flor Tre35 78.0 95.7. — 6.1
113 64) 500) B57 88 104.4 TOA) LON 237 76 88 102.7 86.0 04.3 — 7.5
106 FOr eIGs a ntOs GAS Ai oyys) 105 77) 1130) 80 76 90.8 77.0 83.9 —I7.9
94 IIS Mean 101.8
As Bo
Aug. June May Aug. June July Mean Aug. June May Aug. June Mean way Mean
AS a yey ED EE 409)) 23) Wy22aie23) aera
53 99 79 Scmersaunge 97.0 84 74 110 4100 86 90.8 106.2 98.5
80 118 80 88 128 137 105.2 89 62 79 99 81 82.0 98.0 90.0
107 121 78 Sip er2set25) | Loos IOI 77 67) nr 4! 87 89.2 107.2 98.2
IOI 115 90 65 Serr 106 98.0 116 87 60 115 86 93.8) 17563. Sro4ls
138 Q5| TI2 85 Srars2 | 507.2 134 92 86 “107 “113 106.4) | 10057) Vrod:o
Tes LOON menT Ob) (ee LO7a LTO) 6 Ers53 123 95 83 92 “IIT 100:8 109.8 “Tose
124 SO) Ds5 C7 LOM eT 22) se TOOL7, IIo 97 92 95) mrs, | 1024 mors) Samie
05 83 «158 (ofey agri Faure) etoye pte) 116 85 87 Shi | iyAXe) 99.0) 10T-2) Prosar
105 9006 164 126 1027 129 120.8 IOI 75) ‘£00 LOS) 1140), ods2!) \itoGss) wroses
Tie LOA) TS5)) eTOLn e102) ple4 || TTT. 2 58 79 Ot euro) e132 93-8 104.5 99.1
80 79 37,0 85 Sr 7243. 106.3 88 58 93° 50 lirr8) ‘ror-4 Tox hogs
84 907 144 99 69 134 104.5 96 65 o2 Iq OITA O42) Tito T0746
So L0O) RIO) jr22 68e er38i = 107.3 90 79 or 9154 Or ToO3;0}" T1sl0) Proo1o
COT 24 erro met oS 6208154. “I2r-0 98 84 S30) 1rd) ErT0 97.8 97.0 97.4
04.133) 103), 146 WE Ego! | LI5.0 100 70 96 106 87 91.8 105.2 98.5
110 Mean 102.8
As Bs
Nov. Sept. Nov. Sept. Dec. Mean Nov. Sept. Dec. Dec. Sept. Mean AsYs Mean A
Te eeOO) MeO) sO Sms O 00 = 09 BCOM nn a2Omees3
GW ab 98 97 97 97.4 SAULT Om mr 82 100.4 105.4 102.9 -—+10.6
S2 une 75 102 87 91.6 85 132 97 90 83 97-4 106.8 102.1 +9.8
88 100 63 91 86 85.6 O36) L152) © 200 86 76. (103-25 T0816) rogt4 seen
III 100 59 81 83 86.8 80 149 101 87 55 94.4 109.0 103.2 +10.9
135 90 70 77 79 90.2 S45 307, T7r2 71 50 84.8 102.4 93.1 + 0.8
T718 O7e LOSE OS eee SOME O Oe S55) (O3ir (99) 954. 56) 477240 07.4) ee Ae
159 98 120 72 64 102.6 940 | 90)" 6945)) 53" 1 45E) 73.2) VOTtOn yuis2sqa moro
T5ON OCH 2a C4 ieee O 7am DOSS <4. 73 po) Ni winch tin 50 6: 69.6_—Ss 85.6 776) = Tay
139 80 106 67 77 93.8 81 Ope 020 63 73 84.4 86.8 85.6 — 6.7
127 85 129 72 82 99.0 81 Fou TES 70 Gi) 86.0 90.2 88.1 — 4.2
137 Fou §L27, 83 104 105.4 61 68 123 70) 1103 86.2 96.2 91.2 — II
144 62 136 85 107 106.8 62 58 I14 Cpe seals 84.2 102.6 03-4 +d
138 62 127 TSMNAL 3 eLOS <0 59 61 89 ys LAD 8s. 103.4 94.2 + 1.9
138 59 «6138 85 9 125.) 100:0 60 55 83 64) 223 77.0 93.8 85.4 — 6.9
149 co -LLTy, 79 ILO!) -LO2.4 92 80 Io1 53. p27 90.6 99.0 94.8 + 2.5
Mean 92.3

lines, represents the first half of the interval. Along with it, in heavy
dotted lines (b) appears the same curve, shifted in phase as indicated
in the tabulation, tables 1 and 2. The medium-heavy full curve above
(c) in each chart, gives the results tabulated for the second half of

+++) t+4+44+1 ||

AU OARNO SD ODADAS

Oop & DAUR doth WN DOOD

NO. 16 PRECIPITATION AT PEORIA—ABBOT DE

the interval. The reader will perceive a considerable similarity in
form between curves (b) and (c). Owing to the causes producing
roughness in the mean results, as explained above, this correspondence

oe
eurnua
eT ie

Sang

va
wn
ay

Ao
~
Ys
-
-
-
-
N
N
!
!
D

Fics. 2 (lower) AND 3 (upper).—Graphs of the 97¢-month periodicity.

is not as close as one could wish. However, it is the best available,
and mean curves in heavy lines are plotted in lines (d). Their general
similarity to the curves (c) indicate that the first half of the data,

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

when in the same phase, supports the second-half determinations of
the periodicity.

Amplitudes of Peoria periodicities —The reader will note that all
the curves (d) in figures 2 and 3 indicate an amplitude for the 9-
month periodicity of above 10 percent and several of them above 20
percent. Of the 19 periodicities determined in Peoria precipitation,
none have a less average amplitude than 5 percent, and 10 of them

PAGES HEH
hy My wan [MA _|

Fic. 4.—Graphs of the 154-month periodicity.
Wolf numbers > 20.

exceed 20 percent in average amplitude. This may seem extraordinary
to those who recall that the solar variations which control these perio-
dicities in precipitation, as recorded in the paper cited above, seldom
had amplitudes as great as 0.2 percent. But the explanation is not as
yet susceptible to theory ; we have to accept the results of tabulations.
Figures 4 and 5, suited to Wolf numbers = 20 for the periodicity
of 154 months, will be understood from the preceding description,
relating to 94 months. It will be noted that in figures 4 and 5 the
amplitudes of the curves (d) are in all cases about 20 percent.

NO. 16 PRECIPITATION AT PEORIA—-ABBOT bs

Forecasts of precipitation for future years—tThe reader will per-
ceive that since the periodicities fixed by tabulations of Peoria precipi-
tation employ only data of the years 1856 to 1939, it is perfectly justi-
fied to use them to predict from a synthesis the precipitation expected
to be observed at Peoria in the years 1940 to 1950. This operation
has been performed. The data for it are such as given in the columns A
of tables 1-4.

ood aa ma
PONSA
AWC
CCW CA
el zLAD ed NL

MISA
et
15 ae

Fic. 5.—Graphs of 154-month periodicity.
Wolf numbers < 20.

To fix ideas, there is given in table 5 as much of this tabulation as
covers the years 1940 and 1950. The summation of the 19 columns
purports to show for these future years what will be the percentage
departures from the normals that were printed above. In the next to
final column of table 5 are given the actual observed departures from
those normals, computed from Weather Bureau records. The final
column gives the percentage differences between predicted and ob-
served. The reader will recall that all these data were smoothed by
5-month consecutive means before the tabulation.

nary ean ere

NFMOHMONROA TN
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TONNO NO MOO
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uoynryiaaa pun uoysipasg sisayjuds fo ajguog—sS alav]

weee

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see.

ener

‘29q
“AON
190
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“sny
4in{
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-60 48 1949 1950
d in Smithsonian Publ. 4088.
| 5-month running means.

PRE

+40

MWA NA WR Le A | ly lO YN
bs MAO NS BAVA AA TE

i IN NAINA OSC
mee Wy ie i

acy

194] 1942

‘

| t oe ei ee a et eee” ee ee Seen
1943 V * 1944

Fig. 6.—Predicted precipitation at Peoria, IIl.,

1945 1946 1947 1948 1949
1940 to 1950, and verification. Prediction based on sun’s variation as determined in Smithsonian Publ. 4088
Thin line is prediction, thick line is event. Dotted line is prediction altered in phase as stated in text.

1950
All curves from 5-month running means

NO. 16 PRECIPITATION AT PEORIA—ABBOT 15

Graphical comparison of forecast and event.—In figure 6 the whole
march of the synthetically forecasted expected departures from nor-
mal precipitation at Peoria from 1940 to 1950 are plotted along with
the actual departures observed. The reader will still recall that the
data were smoothed by 5-month running means before any tabulating
was done.

Fit of the curve of prediction —For 3 years, 1940 to 1942, there is
fair agreement between prediction and event. The average divergence
between the two curves for 37 months is 13.9 percent. This is quite
as close an agreement between prediction and synthesis of periodicities
as obtained in the years prior to 1940, when, as one might say, the
curves are “tailored to fit.” Illustrations of this are given in figures 7
and 8, which deal with the intervals 1890 to 1895 and 1911 to 1916.

Phase changes.—After 1942 came changes of phase, but not of
form. Heavy dotted curves are drawn in figure 6 to show the great
similarity of form of the two curves when certain changes of phase
are permitted. From October 1942 to July 1943 the predicted curve
is moved forward 6 months. From March 1944 to June 1945 the
predicted curve is moved backward 3 months. Thereafter, for the
long interval extending until July 1949, the predicted curve is moved
backward bodily 6 months. Finally, from February 1949 to February
1950 the predicted curve is moved forward 6 months.

With these changes of phase admitted, the two curves show for 81
months after 1942 an average divergence between them of 15 percent.
Adding the first 3 years, which showed 35 months of fairly close fit,
prediction would agree with event for 116 months out of 132, with
an average divergence of 15 percent, though including several large
swings of from 70 to 90 percent in amplitude. With such a good
measure of success an 11-year prediction of precipitation at Peoria
would confront us, though based on solar variation alone with no re-
course to meteorology, if the several phase changes above suggested
could be understood and anticipated.

Taking no consideration whatever of phase changes, the average
departure between prediction and event is 14.7 percent for 99 months
out of 136. Large divergences occur in the other 37 months.

Quality of representation—To throw light, if possible, on phase
changes, and to expose the roughness of the representation of precipi-
tation by syntheses of periodicities as it stands before attempting a
prediction, owing to causes already discussed, two intervals of curves
of synthesis and observation prior to 1940 are shown in figures 7 and 8.
The intervals chosen are from 1890 to 1895 and from 1911 to 1916.
The synthetic curve in figure 7 is at a disadvantage, compared to that

VOL. 117

SMITHSONIAN MISCELLANEOUS COLLECTIONS

*}X9} UT syUSWIUIOD 93g ‘OrOI
0} so1Id sreak Ul JWOAS YPM sisayyUAs jo suostsedwoy— (Joddn) g GNV (JOMOT) 4 ‘SOI

. Pil te hast
PST RANT APE
CORA Ha
esi URE 21
Apert rarer an cere rae
HAA
avy a AAA E LN
ah Se, VATA
A
CORTE

NO. 16 PRECIPITATION AT PEORIA—ABBOT it 7/

in figure 8, because, as stated above, all the mean values from tabula-
tions of the first half of the data were shifted in phase as required to
match the phases of the mean values of data of the second half. Hence
the general mean used for the syntheses is not so well suited to the
first half of the 84-year interval as to the second half.

This disadvantage shows plainly, for figure 7 shows less good agree-
ment than figure 8, yet a considerable part of the curves in figure 7
show fair correspondence. What is particularly interesting in figure 7
is that several cases of phase shifting, similar to those noted above in
figure 5, are obvious. Such shiftings appear in the years 1890, 1892,
1893, and notably in 1895. One clearly marked phase shift is seen in
the year 1913, in figure 8, and a lesser one in the year 1916.

The reader will see, by comparing figures 7 and 8, which relate to
years employed for computing the data for prediction, with figure 6,
where the basic periodicities were used for forecasting, that the
“tailored to fit” curves, figures 7 and 8, agree no better, if indeed quite
as well, with observation as the predicted curve agrees with the event
in figure 6.

Solar variation controls weather—The outcome of this attempt to
forecast precipitation for a decade in advance by knowledge of perio-
dicities in solar variation, without recourse to meteorology, is some-
what disappointing because of the phase changes encountered. It had
been hoped that forecasts of percentage precipitation for coming
seasons Over a 10-year interval to within 10 or 15 percent might be
made. If this could be done for one station, perhaps it might also be
done for many. Then a network of lines of equal percentage precipita-
tion for coming seasons for several years in advance might be laid
down on the map. Possibly meteorologists, if they take interest in the
idea, may find some means to conquer the phase-changing obstacle,
and realize this dream of long-range seasonal forecasts.

Hitherto, for 40 years there has been a reluctance on the part of
scientists, and especially meteorologists, to recognize the reality of
observed changes of solar radiation. Even if it were admitted that
very small solar changes occur, they were thought by these scientists
to be insignificant for weather. As one said: “If a room be lighted
with 100 lamps, and one is extinguished, no one could notice the loss
of illumination.” This paper, and Smithsonian Publ. 4088, indicate
a contrary conclusion.

SUMMARY

I have sought, in a preceding paper cited above (footnote 1) to
demonstrate the reality of 23 periodic changes in the intensity of the

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

solar rays that warm the earth. In this present paper it is shown that
periodicities of the same length as 19 of those discovered in solar
variation are accompanied by changes of 5 to 20 percent amplitude in
the precipitation at Peoria, Ill. By samples graphed in figures 2, 3, 4,
5, and in no way unrepresentative of all the periodicities tabulated, it
is shown that these periodic variations of precipitation occur in nearly
equal amplitudes and with moderate differences of phase in 12 separate
independent intervals, each several years long, between 1856 and 1939.

Using these results as a basis, I predict the precipitation for Peoria
from 1940 to 1950. With a range of actual precipitation from 50 to
150 percent of normal, occurring in that interval in the smoothed
records, prediction matches the event for 8 out of 11 years to within
an average deviation of 14.7 percent. In the other 3 years, large fluc-
tuations in actual precipitation were matched by strikingly similar
large fluctuations in the curve of prediction, but with phase differences
of from 3 to 6 months. The prediction matches the event as well after
Io years as at the beginning. The fluctuations in the curve of predic-
tion have almost precisely the same scale of range as those in the curve
of actual precipitation.

While it is difficult to conceive, theoretically, how solar changes,
seldom as large as 0.2 percent, could produce variations of from 5 to
20 percent in precipitation, the facts are there. It may be that adequate
theory will eventually be found to explain them. In the meantime it
would be of no importance, practically, whether theory had overtaken
fact, if it were found that these periodic changes of precipitation could
be synthesized to forecast seasons for years in advance.

A partial success in that direction has been achieved. For 3 years
in advance the curve of prediction shown in figure 6 follows generally
a rise of actual precipitation amounting to 90 percent. For 99 months
out of 136, indeed, the average divergence is 15 percent. But the
long-range prediction is marred by changes of phase, which, thus far
at least, cannot be anticipated and allowed for in advance.

To the interested general reader, unfamiliar perhaps with con-
sidering percentages of normal precipitation, the meat of the matter
may be expressed as follows: A prediction for 11 years in advance,
based on knowledge of the sun’s variation, without any consideration
of meteorology, fits the rainfall curve at Peoria, IIl., for 99 months out
of 136, to within an average error of one medium rainfall per month,
or about 0.4 of an inch per month. Sometimes, however, dry or wet
spells occur 3 to 6 months from when they are expected. Unless these
phase changes can be anticipated, the method fails to come up fully to
what is hoped for.

_ SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 17

A GENERIC SYNOPSIS OF THE LIZARDS
OF THE SUBFAMILY LYGOSOMINAE

BY
M. B. MITTLEMAN

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NOVEMBER 4, 1952

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 117, NUMBER 17

meen SYNOPSIS OF THE LIZARDS
OF THE SUBFAMILY LYGOSOMINAE

BY
M. B. MITTLEMAN

(Pusuication 4096)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION

NOVEMBER 4, 1952

The Lord Baltimore Press

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A GENERIC SYNOPSIS OF THE LIZARDS OF
THE SUBFAMILY LYGOSOMINAE?
By M. B. MITTLEMAN

As long ago as 1887 Boulenger remarked on the difficulty he had
encountered in attempting to make a satisfactory generic arrange-
ment of the Scincidae. In the long interval from Boulenger’s day to
our own, the skinks have reaffirmed their characteristic refractori-
ness to systematic order, and by far the most notorious offenders
have been those groups of which the genus Lygosoma and certain of
its allies are representative. These lizards, known loosely as the
“lygosome” genera, are as baffling to the taxonomist as they are
diverse in the multiplicity of their forms. Probably not the least
reason for the perplexity surrounding the lygosomes is the fact that
they add or delete many supposedly fundamental characters with the
most disconcerting irregularity, while at the same time presumably
trivial traits are often maintained through great arrays of species
which are distributed over large areas.

The Boulengerian arrangement of genera was based on the recog-
nition of these supraspecific groups according to the presence, absence,
or other variation of certain characters accorded “generic” impor-
tance. Such a system inevitably becomes unwieldy when applied to
lizards displaying the attributes of the lygosomes and results also in
some weird zoogeographic and phylogenetic conclusions. On the
other hand, in defense of Boulenger’s arrangements and the emenda-
tions suggested by M. A. Smith (1937), the recognition of a rela-
tively few genera considerably eases the every-day classification of the

1For many helpful suggestions in the preparation of’ this study, as well as
for the opportunity to examine specimens in his care, I am grateful to Arthur
Loveridge, of the Museum of Comparative Zoology. To Dr. Hobart M. Smith,
of the University of Illinois, I wish to express my appreciation for the loan
of literature, for checking references, and for numerous friendly and construc-
tive criticisms. Finally, I must acknowledge my considerable obligation to the
authorities of the Smithsonian Institution for the generous use of facilities at
the United States National Museum and for the publication of this paper; and,
particularly, to Dr. Doris M. Cochran, who has generously cooperated with me
in innumerable ways over the course of several years, has given the present
manuscript a careful reading, and has lent me specimens in her care.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 17

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

lygosomes, and if the ultimate purpose is merely to pigeonhole a
particular species at hand, the Boulengerian approach is certainly as
good as any other.

The question then arises as to whether some other arrangement is
desirable for the lygosome genera and, if so, upon what basis such a
system should be predicated. The obvious antithesis of Boulenger’s
taxonomy is one wherein genera result from the “brigading of
demonstrably related species,” and where more tightly drawn generic
definitions are made. It seems to me that this latter system is in-
comparably superior to that of Boulenger, for it has the dual ad-
vantages of reflecting phylogenetic relationships and of restricting
the exercise of arbitrary judgment of characters deemed to be of
“generic” importance. The necessity for some such arrangement
goes far beyond the private philosophies of the “lumpers” and the
“splitters” if any kind of comprehensible and stable arrangement is
to be brought to the more than 600 described forms of lygosome
lizards.

Since the most recent review of the lygosomes is that of M. A.
Smith (op. cit.), reference to and comparison with this author’s
arrangement of these lizards are necessary. In the summary that fol-
lows it will be seen that the supraspecific arrangement suggested here
differs from that of Smith in two basic ways: First, that the lygosome
genera are regarded as a natural subfamily of the Scincidae, and
second, that genera alone are utilized instead of genera, subgenera,
and sections. In arriving at the conclusions presented here with re-
spect to genera I have employed the following guiding criteria: First,
that genera should be aggregations of species sharing one or more
constant traits which set them apart from all other species groups;
second, that genera should consist of species of common phylogenetic
origin as far as this may appear determinable; third, that, in general,
compact and more tightly drawn genera are preferable to loosely de-
fined, all-inclusive groupings. Granting that my generic criteria are
probably as personal (and therefore as vulnerable) as may be those
of M. A. Smith, I believe they are defensible to the extent that they
are essentially consistent and that they indicate relationships, and
beyond this I think the genus cannot reasonably be expected to go.
In brief, the 600-odd described forms of lygosome lizards individually
and collectively constitute the best possible argument for the discrimi-
nation of discrete generic groups which are refined beyond the stage
of catch-all depositories for widely divergent forms of uncertain
origin.

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 3

The essence of the arrangement used here, and the differences
between this system and that of Boulenger, as emended by M. A.
Smith, are as follows:

1. Lygosoma and certain related genera are recognized here as
constituting a subfamily (Lygosominae) of the Scincidae, which is
further divided into three additional subfamilies. Smith does not
divide the Scincidae into subfamilies.

2. The subfamily Lygosominae is regarded as consisting of at
least 33 genera. Smith recognized 10 genera, 3 sections, and 5 sub-
genera of Lygosoma and its allies.

3. The following lygosome genera are recognized:

Ablepharus Fitzinger, 1823.

Anotis Bavay, 1860.

Ateuchosaurus Gray, 1845.

Carlia Gray, 1845.

Cophoscincopus Mertens, 1934.
Cophoscincus Peters, 1867.
Cryptoblepharus Wiegmann, 1834.
Dasia Gray, 1839.

Emoia Gray, 1845.

Eugongylus Fitzinger, 1843.

Eumecia Bocage, 1870.

Hemiergis Wagler, 1830.
Lampropholis Fitzinger, 1843.
Leiolopisma Duméril and Bibron, 1839.
Leptosiaphos Schmidt, 1943.

Lipinia Gray, 1845.

Lygosoma Hardwicke and Gray, 1827.
Mochlus Giinther, 1864.

Nodorha, new genus.

Norbea Gray, 1845.

Ophioscincus Peters, 1873.

Otosaurus Gray, 1845.

Panaspis Cope, 1868.

Rhodona Gray, 1839.

Riopa Gray, 18309.

Ristella Gray, 1839.

Saiphos Gray, 1831.

Scincella Mittleman, 1950.
Sphenomorphus Fitzinger, 1843.
Squamicilia, new genus.

Tachygyia, new genus.

Tribolonotus Duméril and Bibron, 1839.
Tropidophorus Duméril and Bibron, 1839.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 107

4. The supraspecific groups recognized by M. A. Smith are as
follows:

Genus:

Ablepharus Fitzinger, 1823.
Ateuchosaurus Gray, 1845.
Cophoscincopus Mertens, 1934.

Dasia Gray, 1830.

Emoia Gray, 1845.

Lygosoma Hardwicke and Gray, 1827.
Otosaurus Gray, 1845.

Rhodona Gray, 1839.

Riopa Gray, 1830.

Tiliqua Gray, 1825.

Subgenus :

Eugongylus Fitzinger, 1843.
Eumecia Bocage, 1870.
Ictiscincus M. A. Smith, 1937.
Panaspis Cope, 1868.

Riopa Gray, 18309.

Section:

Leiolopisma Duméril and Bibron, 18309.
Lygosoma Hardwicke and Gray, 1827.
Sphenomorphus Fitzinger, 1843.

5. The following groups used by M. A. Smith are not recognized
here, for the reasons given:

Ictiscincus M. A. Smith, 1937—Sphenomorphus Fitzinger, 1843 (vide
Loveridge, 1948, p. 352, regarding the supposed distinguishing charac-
teristics of the teeth).

Tiliqua Gray, 1825. Not a lygosome genus (see following discussion and
vide Waite, 1920, pp. 132-133).

The 33 genera previously enumerated constitute what I conceive
to be the subfamily Lygosominae of the family Scincidae. The char-
acteristics of the Lygosominae and of the three other subfamilies
of the Scincidae are summarized in the following synopsis:

A. Palatine bones in contact on median line of palate.

I. Pterygoid bones separated on the median line of palate;

palatal notch extending anteriorly to level of centers of

CVS eer SRa a crates “are Mabeve vers ie aie rere eee ite te a erere aaa leneee rete Tete MABUYINAE
2. Pterygoid bones in contact anteriorly; palatal notch not

extending anteriorly to level of centers of eyes.......... LyGOSOMINAE

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 5

B. Palatine bones separated on median line of palate.

1. Nostril pierced in nasal, or between two adjacent plates, but

Mever tOMeMne” FOStral se. 2. an dca ut ceo SCINCINAE
2. Nostril pierced between rostral and an adjacent plate,

thereby contacting rostral, or else within rostral itself..CHALCIDINAE

The Mabuyinae are distributed in Indo-Malaya, Africa, Mada-
gascar, Asia, and the Americas (including the West Indies). The
Lygosominae have an approximately similar distribution but occur
also in southern Europe, Australasia, and the Pacific Islands. The
Scincinae virtually duplicate the Lygosominae in their occurrence,
while the Chalcidinae are restricted to Africa, Madagascar, southern
Asia, and southern Europe.

The general plan of organization in this paper incorporates a brief
diagnosis for each lygosome genus, or else the distinguishing features
which set apart the genus from other lygosome genera are enumer-
ated. Included also are the genotype for each genus and the principal
synonyms. In addition to an artificial key to the genera, there are
appended alphabetical listings of primary generic synonyms and
species generically allocated according to the taxonomy suggested
here.

Genus LYGOSOMA Hardwicke and Gray

Lygosoma HarpwickeE and Gray, Zool. Journ., vol. 3, p. 228, 1827. (Type:
Lacerta serpens Bloch= Anguis quadrupes Linné.)

Podophis WiEGMANN, Herpetologia Mexicana, p. 11, 1834. (Type: Anguis
quadrupes Linné.)

Eyelids well developed, movable, the lower one scaly and lacking
a more or less translucent or transparent disc; no supranasals ; pre-
frontals small and widely separated; frontoparietals united, distinct
from the large interparietal; preanals not, or barely, enlarged; ear
opening absent or punctiform, approximating nostril or smaller if
present ; limbs much reduced, digits 5-5 ; body elongate, slender.

Distribution.—Australia, Java, Malaya, Siam, Indo-China, Philip-
pine Islands.

Genus SPHENOMORPHUS Fitzinger

Eulamprus Frtzincer, Systema reptilium ... , p. 22, 1843. (Type: Lygosoma
quoyt Duméril and Bibron.)

Sphenomorphus Frrz1ncEr, ibid., p. 23. (Type: Lygosoma melanopogon Duméril
and Bibron.)

Hinulia Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 22, 1845. (Type: Lygosoma quoyi Duméril and Bibron.)

Elania Gray, ibid., p. 80. (Type: Scincus muellert Schlegel; non Elania
Sundevall, 1835.)

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOs Thy,

Lissonota Biytu, Journ. Asiatic Soc. Bengal, vol. 22, p. 653, 1853. (Type:
Lissonota maculata Blyth; non Lissonota Gravenhorst, 18209.)

Ictiscincus M. A. SmitH, Rec. Indian Mus., vol. 39, No. 3, p. 222, 1937. (Type:
Scincus muellert Schlegel.)

Differs essentially from Lygosoma as follows: Prefrontals large,
often forming a median suture; frontoparietals paired; preanals en-
larged (except in hallieri) ; ear opening always present, large, some-
what smaller than the eye; limbs large and separated slightly or
meeting or overlapping when appressed.

Distribution Australia, East Indies, New Guinea, India, Indo-
China, Malaya.

Note.—This is by far the largest lygosome genus in point of in-
cluded forms, well over 150 having been described. Quite possibly
the genus as here constituted is polyphyletic, but until considerably
more information can be accumulated on the individual and popula-
tion variations of these lizards they are best considered as congeneric
despite the fact that certain species groups may be worthy of generic
rank.

Genus SAIPHOS Gray
Saiphos Gray, in Griffith's Animal Kingdom, Syn. 9, p. 72, 1831. (Type: Seps
equalis Gray.)
Peromeles WIEGMANN, Herpetologia Mexicana, p. 11, 1834. (Type: Seps
equalis Gray.)
Anomalopus DuMERIL, Catalogue méthodique de la collection des reptiles... ,
p. 185, 1851. (Type: Anomalopus verreauxii Dumeéril.)

Coloscincus PETERS, Monatsb. Akad. Wiss. Berlin, 1876, p. 532. (Type:
Coloscincus truncatus Peters.)

Differs essentially from Lygosoma as follows: Prefrontals greatly
reduced, or often absent ; frontoparietals paired; preanals enlarged ;
ear opening absent; limbs minute, reduced to tridactyle, didactyle, or
monodactyle rudiments.

Distribution —Australia, Sumatra, Malaya.

Genus LEPTOSIAPHOS Schmidt

Leptosiaphos K. P. Scuint, Zool. Ser. Field Mus. Nat. Hist., vol. 24, No. 29,
Pp. 332, 1943. (Type: Lygosoma meleagris Boulenger.)

Differs essentially from Lygosoma as follows: Frontoparietals
paired ; limbs well developed, digits 3, 4, or 5.
Distribution —East Africa.

Genus OTOSAURUS Gray

Otosaurus Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 93, 1845. (Type: Otosaurus cumingi Gray.)

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN Wi

Parotosaurus BouLencer, Trans. Zool. Soc. London, vol. 20, p. 257, 1924.
(Type: Parotosaurus annectens Boulenger.)

Differs essentially from Lygosoma as follows: Supranasals usu-
ally present, either small and widely separated or else well developed
and forming a median suture; frontoparietals paired; preanals en-
larged ; ear opening large, approximating eye; limbs large, separated
slightly or meeting or overlapping when appressed.

Distribution—Kast Indies, Borneo, Sarawak, Sumatra, Malaya,
New Guinea, Philippine Islands.

Note—The genus Parotosaurus may be used conveniently for
those forms possessing reduced nonsuturing supranasals, or else lack-
ing these scales; in this case Otosaurus would then be restricted to
the species with enlarged supranasals which form a median suture.

Genus COPHOSCINCUS Peters

Cophoscincus Peters, Monatsb. Akad. Wiss. Berlin, 1867, p. 19. (Type: Lygo-
soma (Cophoscincus) quadrivittatus Peters.)

Differs essentially from Lygosoma as follows: Preanals enlarged ;
ear opening absent ; limbs robust and either separated by several scale
lengths when appressed or else meeting or overlapping.

Distribution—Australia, Sumatra, Borneo, Malaya, Celebes, Philip-
pine Islands.

Note.—In quadrivittatus the lower eyelid is scaly and lacks a trans-
parent or translucent disc, while in other species (cf. infralineolatus,
relictus) the disc is present. In this group the presence or absence
of the palpebral disc is not generically important.

Genus DASIA Gray

Dasia Gray, Ann. Mag. Nat. Hist., vol. 2, p. 331, 1839. (Type: Dasia olivacea
Gray.)

Lamprolepis Frrzincer, Systema reptilium, p. 22, 1843. (Type: Scincus
smaragdinus Lesson.)

Liotropis Firzincrr, ibid., p. 22. (Type: Euprepres ernestii Duméril and
Bibron = Dasia olivacea Gray.)

Keneuxia Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 79, 1845. (Type: Scincus smaragdinus Lesson.)
Apterygodon Eprrttnc, Nat. Tijdschr. Med. Ind., vol. 26, p. 483, 1863. (Type:

Apterygodon vittatum Ederling.)
Theconyx ANNANDALE, Spolia Zeylanica, vol. 3, p. 191, 1906. (Type: Euprepes
halianus Haly and Nevill; non Theconyx Gray, 1845.)

Differs from Lygosoma essentially as follows: Supranasals present,
occasionally forming a median suture, sometimes partially or nearly
completely fused with the nasals; prefrontals large, sometimes form-

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

ing a median suture; frontoparietals paired; body and limbs robust,
the latter often overlapping when appressed, or else separated by only
a few scale lengths.

Distribution Indian Archipelago, Papuasia, Caroline and Philip-
pine Islands, India, Borneo.

Genus COPHOSCINCOPUS Mertens

Cophoscincopus MERTENS, Zoologica (Stuttgart), vol. 32, p. 190, 1934. (Type:
Cophoscincus simulans Vaillant; non Cophoscincus Peters, 1867 = Tiliqua
dura Cope.)

Differs from Lygosoma essentially as follows: Supranasals present
and forming a median suture; frontoparietals paired; ear opening
present but almost completely covered by overlapping scales ; preanals
enlarged ; limbs robust and long, overlapping when appressed.

Distribution—West Africa.

Genus ATEUCHOSAURUS Gray

Ateuchosaurus Gray, Catalogue of the specimens of lizards in the collection of
the British Museum, p. 107, 1845. (Type: Ateuchosaurus chinensis Gray.)

Lygosaurus HatitowELt, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 406.
(Type: Lygosaurus pellopleurus Hallowell.)

Differs essentially from Lygosoma as follows: Frontoparietals
paired ; frontal long and constricted (longer than frontoparietals and
interparietal together) ; parietals much reduced; ear opening large,
about midway in size between eye and nostril; limbs robust, either
overlapping when appressed or separated by a few scale lengths.

Distribution—China, Indo-China, Ryukyu Archipelago.

Genus RIOPA Gray

Riopa Gray, Ann. Mag. Nat. Hist., vol. 2, p. 332, 1839. (Type: Lygosoma
punctata Gray = Lacerta punctata Linné.)

Chiamela Gray, ibid., p. 332. (Type: Chiamela lineata Gray.)

Hagria Gray, ibid., p. 333. (Type: Hagria vosmaerii Gray.)

Campsodactylus DuMERIL and Bipron, Erpétologie générale . . . , vol. 5, p. 761,
1839. (Type: Campsodactylus lamarrei Duméril and Bibron=Hagria
vosmaerit Gray; non Campsodactylus Duméril, 1837, nom. nud.)

Sphenosoma FItzincEr, Systema reptilium, p. 23, 1843. (Type: Ewumeces
punctatus Wiegmann — Lacerta punctata Linné.)

Eyelids well developed, movable, the lower one with a more or less
transparent or translucent disc; supranasals present, forming a
median suture; prefrontals small, widely separated; frontoparietals
paired, distinct from interparietal; preanals not, or barely, enlarged ;

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 9

ear opening small (punctiform), approximating nostril or smaller ;
limbs much reduced, digits 5-5 or less; general habitus vermiform.
Distribution—India, Burma, Kenya.

Genus EUGONGYLUS Fitzinger

Eugongylus Fitzincer, Systema reptilium, p. 23, 1843. (Type: Eumeces
oppellii Duméril and Bibron = Lacerta rufescens Shaw (part).)

Differs from Ritopa essentially as follows: Lower eyelid either
uniformly scaly or else with a small opaque disc ; supranasals reduced
and not forming a median suture; limbs well developed, digits 5-5;
body robust, lacertiform.

Distribution —Australia, Papuasia, Halmahera, Sula Islands.

SQUAMICILIA, new genus

Type: Eumeces isodactylus Giinther, The reptiles of British India,
p. 93; pl. 13; fig. A, 1864.

Differs essentially from Riopa as follows: Lower eyelid scaly, lack-
ing a more or less transparent or translucent disc; supranasals re-
duced and not forming a median suture; frontoparietals usually
united, interparietal much reduced; ear opening reduced, often only
a small oblique slit.

Distribution— Siam and Somaliland.

Genus MOCHLUS Giinther

Mochlus Gintuer, Proc. Zool. Soc. London, 1864, p. 308. (Type: Mochlus
punctatus Giinther—=Eumeces (Riopa) sundevallii Smith.)

Sepacontias GUNTHER, Ann. Mag. Nat. Hist., ser. 5, vol. 6, p. 235, 1880. (Type:
Sepacontias modestus Giinther.)

Lepidothyris Corr, Journ. Morph., vol. 7, p. 233, 1892. (Type (by subsequent
designation) : Lepidothyris fernandi Cope = Tiliqua fernandi Burton.?)

Differs essentially from Riopa as follows: Lower eyelid scaly, lack-
‘ing a more or less transparent or translucent disc; limbs robust,

2 Cope’s original description of Lepidothyris (loc. cit.) is merely that of a
scincid genus of lizards with no included species, and appears only in a key to
the genera of Scincidae. In accordance with Opinion 46 of the International
Commission “the first species published in connection with the genus . . . be-
comes ipso facto the type.” Thus the type of Lepidothyris is the species
fernandi, since Cope (1900, p. 617) mentions Lepidothyris fernandi briefly. De-
scribed with Lepidothyris at the same time and in the same fashion are the
following genera: Monophorus, Oncopus, Ollochirus, Tridentulus, Furcillus,
Haploscincus, and Dimeropus. All these are scincid genera; presumably Lygo-
sominae, with no included species, and must at present be considered unidenti-
fiable nomina inquirendae, except for Monophorus which is clearly unavailable
since it is preoccupied by Monophorus Deshayes, 1877 (Mollusca).

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

barely meeting or else separated by several scale lengths when ap-
pressed ; digits 5-5.

Distribution—Africa, China, Indo-China, Malaya, Philippine
Islands.

TACHYGYIA, new genus

Liosoma FitzincEer, Systema reptilium, p. 22, 1843. (Type: Eumeces micro-
lepis Duméril and Bibron; non Liosoma Brandt, 1834.)

Type: Eumeces microlepis Duméril and Bibron, Erpétologie
générale, vol. 5, p. 659, 1839.

Differs essentially from Riopa as follows: Lower eyelid scaly, lack-
ing a more or less transparent or translucent disc; frontoparietals
united, interparietal reduced; limbs very robust and long, broadly
overlapping when appressed.

Distribution —Friendly Islands, New Caledonia.

Genus EUMECIA Bocage

Eumecia Bocace, Journ. Acad. Sci. Lisbon, vol. 3, p. 67, 1870. (Type: Eumecia
anchietae Bocage.)

Differs from Riopa essentially as follows: Prefrontals large, form-
ing a median suture ; ear opening large (only slightly smaller than the
eye) ; limbs minute; digits 2-3 (in Riopa 3-3 or more).

Distribution—British East Africa, Nyasaland.

Genus PANASPIS Cope

Panaspis Core, Proc. Acad. Nat. Sci. Philadelphia, 1868, p. 317. (Type:
Panaspis aeneus Cope.)

Differs essentially from Riopa as follows: Supranasals reduced, not
forming a median suture, occasionally partially fused with nasals ; ear
opening large, approximating eye; limbs short or long, but robust.

Distribution.—West Africa.

Note.—The species reichenovei and africanus are referred to Leio-
lopisma, although M. A. Smith considers them congeneric with aeneus,
cabindae, and other West African forms which I regard as members
of Panaspis. The habitus is such in reichenovet and africanus as to
render these species generically indistinguishable from many forms
in the genus Leiolopisma. However drastic and improbable the zoo-
geographic implications may be as a result of placing the West African
species in a genus which is largely Pacific in distribution, they are
paralleled by other instances of disjunct distribution in the lygosome
genera, cf. Riopa, Poria, Ophioscincus, Ablepharus, etc.

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN II

Genus EMOIA Gray

Eusoma Fitzincer, Systema reptilium, p. 22, 1843. (Type: Eumeces lessonii
Duméril and Bibron=Scincus cyanurus Lesson; non Eusoma Germar,
1817.)

Emoia Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 95, 1845. (Type: Scincus atrocostatus Lesson.)

Differs from Riopa essentially as follows: Supranasals reduced, not
forming a median suture; interparietal often fused with the single
frontoparietal ; limbs robust and long, overlapping when appressed,
digits 5-5; general habitus lacertiform.

Distribution —Borneo, Philippine Islands, New Guinea, Solomon
Islands, New Hebrides, Fiji Islands, Friendly Islands, Samoa, Ad-
miralty Islands.

Genus LEIOLOPISMA Duméril and Bibron

Leiolopisma DuMERIL and Brpron, Erpétologie générale ..., vol. 5, p. 742,
1839. (Type: Scincus telfairii Desjardins.)

Oligosoma Grrarp, Proc. Acad. Nat. Sci. Philadelphia, 1857, p. 196. (Type:
Mocoa zeylandica Gray — Lygosoma moco Duméril and Bibron.)

Hombronia Gtrarp, ibid., p. 196. (Type: Hombronia fasciolaris Girard.)

Lygosomella Grrarp, ibid., p. 196. (Type: Lygosomella aestuosa Girard.)

Cyclodina Girard, ibid., p. 195. (Type: Cyclodina aenea Girard.)

Eyelids well developed, movable, the lower one with a more or
less transparent or translucent disc; no supranasals; prefrontals
small, separated; frontoparietals paired, distinct from interparietal ;
preanals not, or barely, enlarged; ear opening rather large, often ap-
proximating the eye, always larger than the nostril; limbs well de-
veloped, broadly overlapping or meeting or slightly separated when
appressed, digits 5-5; general habitus lacertiform.

Distribution —Australia, New Caledonia, New Zealand, Tasmania,
Mauritia, West Africa.

Genus LAMPROPHOLIS Fitzinger

Lampropholis Firzincer, Systema reptilium, p. 22, 1843. (Type: Lygosoma
guichenott Duméril and Bibron.)

Eulepis Firzincer, ibid., p. 22. (Type: Lygosoma duperryi Dumeéril and
Bibron = Tiliqua trilineata Gray.)

Mocoa Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 80, 1845. (Type: Lygosoma guichenoti Dumeéril and
Bibron.)

Tropidoscincus Bocacr, Journ. Acad. Sci. Lisbon, vol. 15, p. 230, 1873. (Type:
Tropidoscincus aubrianus Bocage.)

Lioscincus Bocace, ibid., p. 328. (Type Lioscincus steindachnerti Bocage = Lygo-
soma tricolor Bavay.)

Sauroscincus Peters, Sitzb. Ges. Nat. Freunde Berlin, 1878, p. 149. (Type:
Sauroscincus braconnieri Peters—Tropidolepisma variabilis Bavay.)

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

Differs from Leiolopisma essentially as follows: A single fronto-
parietal, well developed, distinct from interparietal; ear opening
moderate (smaller than the eye) to small (approximating nostril or
slightly larger).

Distribution—New Caledonia, Australia, Tasmania, Loyalty
Islands, New Hebrides Islands.

Genus CARLIA Gray

Heteropus DumMériL and Brpron, Erpétologie générale ..., vol. 5, p. 757,
1839. (Type: Heteropus fuscus Duméril and Bibron; non Heteropus
Beauvois, 1805, or Heteropus Fitzinger, 1826.)

Carlia Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 271, 1845. (Type: Carlia melanopogon Gray; non
Lygosoma melanopogon Duméril and Bibron = Sphenomorphus melanopogon
Dumeéril and Bibron, part.)

Myophila pE Vis, Proc. Roy. Soc. Queensland, vol. 1, p. 77, 1884. (Type:
Myophila vivax de Vis.)

Lygisaurus DE Vis, ibid., p. 77. (Type: Lygisaurus foliorum de Vis.)

Differs essentially from Leiolopisma as follows: A single well-de-
veloped frontoparietal; interparietal usually very small or absent;
digits 4-5.

Distribution.—Moluccas, New Guinea, Australia, Papuasia.

Genus ANOTIS Bavay

Anotis Bavay, Mem. Soc. Linn. Normandie, vol. 4, No. 5, p. 29, 1869. (Type:
Anotis mariae Bavay.)

Nannoscincus GUNTHER, Ann. Mag. Nat. Hist., ser. 4, vol. 10, p. 421, 1872.
(Type: Nannoscincus fuscus Ginther = Anotis mariae Bavay.)

Differs essentially from Letolopisma as follows: Prefrontals minute
and widely separated (well developed in Leiolopisma) ; ear opening
absent or punctiform ; limbs much reduced, digits 5-5 or less; general
habitus vermiform.

Distribution—New Caledonia, Australia.

Genus HEMIERGIS Wagler

Tridactylus Cuvier, Régne animal, p. 64, 1829. (Type: Zygnis decresiensis
Fitzinger; non Tridactylus Lacépéde, 1799.)

Tetradactylus Cuvier, ibid., p. 64. (Type: Zygnis decresiensis Fitzinger; non
Tetradactylus Merrem, 1820.)

Hemiergis Wacter, Natiirliches System der Amphibien..., p. 160, 1830.
(Type: Zygnis decresiensis Fitzinger.)

Peromelis Wacter, ibid., p. 160. (Type: Zygnis decresiensis Fitzinger.)

Chelomeles DumMERIL and Brpron, Erpétologie générale ..., vol. 5, p. 774,
1839. (Type: Chelomeles quadrilineatus Duméril and Bibron.)

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 13

Differs essentially from Leiolopisma as follows: Preanals enlarged ;
ear opening minute or absent ; limbs greatly reduced, digits 4-4 or less;
general habitus vermiform.

Distribution —Australia.

Genus LIPINIA Gray

Lipinia Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. 84, 1845. (Type: Lipinia pulchella Gray.)

Differs essentially from Leiolopisma as follows: Prefrontals large,
forming a median suture; frontonasal at least as long as it is wide
(wider than long in Letolopisma) ; preanals prominently enlarged.

Distribution.—Solomon Islands, New Guinea, Philippines, Borneo,
Siam, Tenasserim, Malaya, Andaman Islands.

Genus SCINCELLA Mittleman

Scincella M1TTLEMAN, Herpetologica, vol. 6, No. 2, p. 19, 1950. Type: Scincus
lateralis Say.)

Differs essentially from Leiolopisma as follows: Prefrontals large,
often forming a median suture; preanals prominently enlarged.

Distribution —India, Sarawak, Borneo, North and Middle America,
China, Anadaman Islands, Nicobar Islands, Celebes, Hawaiian
Islands, Tasmania, Australia.

Genus RHODONA Gray

Rhodona Gray, Ann. Mag. Nat. Hist., vol. 2, p. 335, 1839. (Type: Rhodona
punctata Gray.)

Soridia Gray, ibid., p. 335. (Type: Soridia lineata Gray.)

Brachystopus Dumérit and Breron, Erpétologie générale, vol. 5, p. 778, 1839.
(Type: Brachystopus lineopunctulatus Duméril and Bibron=hodona
punctata Gray.)

Praepeditus Dumérit and Brsron, ibid., p. 787. (Type: Soridia lineata Gray.)

Ronia Gray, in Grey’s Travels in Australia, vol. 2, p. 437, 1841. (Type: Ronia
catenulata Gray =Rhodona punctata Gray.)

Pholeophilus A. Smiru, Illustrations of the zoology of South Africa, p. 15
(App.), 1849. (Type: Pholeophilus capensis Smith = Soridia lineata Gray.)

Eyelids well developed, movable, the lower one with a more or less
transparent or translucent disc; no supranasals; prefrontals absent,
or if present then small and widely separated ; frontoparietal single,
fused with the interparietal into a single large shield; preanals en-
larged; ear opening absent or minute; nasals very large and forming
a median suture, or else barely separated; anterior limbs much re-
duced or absent, digits of hind limbs 2 or less; general habitus
vermiform.

Distribution —Australia.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

NODORHA, new genus

Leptosoma Fitzincer, Systema reptilium, p. 23, 1843. (Type: Riopa bougain-
villii Duméril and Bibron (non Leptosoma Leach, 1819).)

Type: Riopa bougainvillii Duméril and Bibron.

Differs essentially from Rhodona as follows: Frontoparietal single
or paired, but always distinct from the interparietal; anterior limbs
present (except in N. wilkinst), digits 5-5 or less.

Distribution.—Australia, Tasmania.

Genus OPHIOSCINCUS Peters

Ophioscincus Peters, Monatsb. Akad. Wiss. Berlin, 1873, p. 747. (Type:
Ophioscincus australis Peters.)

Isopachys LONNBERG, Kungl. Svenska Vet.-Akad. Handl., vol. 55, No. 4, p. 10,
1916. (Type: Isopachys gyldenstolpei Lonnberg.)

Typhloseps ANGEL, Bull. Mus. Hist. Nat. Paris, 1920, p. 4. (Type: Typhloseps
roulet Angel.)

Differs essentially from Rhodona as follows: Lower eyelid immovy-
able, with a transparent disc, or else in itself largely transparent ; no
upper eyelid; frontoparietals paired, distinct from the interparietal ;
limbs absent.

Distribution —Siam and Australia.

Genus ABLEPHARUS Fitzinger

Ablepharus Firzincer, in Lichtenstein, Verzeichniss der Doubletten des zoo-
logischen Museums... Berlin ..., 1823, p. 103. (Type: Ablepharus
pannonicus Fitzinger.)

Lerista Brett, Proc. Zool. Soc. London, 1833, p. 99. (Type: Lerista lineata
Bell.)

Microblepharus Fitz1ncEr, Systema reptilium, p. 23, 1843. (Type: Ablepharus
menestriestt Duméril and Bibron.)

Ophiopsis FirzinceEr, ibid., p. 23. (Type: Levista lineata Bell.)

Menetia Gray, The zoology of the voyage of H.M.S. Erebus and Terror... ,
Rep., pl. 5, fig. 4, 1844. (Type: Menetia greyi Gray.)

Miculia Gray, ibid., pl. 5, fig. 3. (Type: Miculia elegans Gray.)

Blepharosteres StoticzKa, Proc. Asiatic Soc. Bengal, 1872, p. 74. (Type:
Blepharosteres grayanus Stoliczka.)

Phaneropus Fiscuer, Arch. Naturg., 1881, p. 236. (Type: Phaneropus muelleri
Fischer. )

Eyelids immovable, a transparent disc covering the entire eye;
supranasals present or absent; frontoparietal single or paired, but
always distinct from the interparietal ; ear opening absent, or if pres-
ent quite small; limbs short but well developed; digits 5-5 or less;
general habitus lacertiform.

Distribution—Australia, Africa, southwest Asia, southern Europe.

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 15

Genus CRYPTOBLEPHARUS Wiegmann

Cryptoblepharus WikGMANN, Herpetologia Mexicana, p. 12, 1834. (Type:
Ablepharus poecilipleurus Wiegmann.)

Morethia Gray, The zoology of the voyage of H.M.S. Erebus and Terror...
Rep., p. 4, 1844. (Type: Morethia anomalus Gray.)

Differs essentially from Ablepharus as follows: Frontoparietals
and interparietal fused to form a single large shield; ear opening
present, large, approximating eye in size.

Distribution —Australia, New Guinea, East Indies, Madagascar,
Africa, Polynesia.

Note.—As remarked by M. A. Smith (1935, p. 309), Ablepharus
appears to be of polyphyletic origin. The separation of certain species
under the generic designation Crypoblepharus primarily takes cog-
nizance of slightly more apparent distinctions than many others which
occur in Ablepharus (sensu lato). It is possible that other species
groups in Ablepharus are generically distinct.

Genus RISTELLA Gray

Ristella Gray, Ann. Mag. Nat. Hist., vol. 2, p. 333, 1830. (Type: JRistella
rurkti Gray.)

Eyelids well developed, movable, the lower one scaly and lacking
a more or less transparent or translucent disc; no supranasals ; pre-
frontals small and well separated, or large and fused into a single
large shield; frontoparietals paired, distinct from the interparietal ;
preanals not enlarged; ear opening well developed; limbs well de-
veloped ; digits 4-5, claws competely retractile into sheaths.
Distribution —Southern India.

Genus TROPIDOPHORUS Duméril and Bibron

Tropidophorus DumériL and Brsron, Erpétologie générale . . . , vol. 5, p. 554,
1839. (Type: Tropidophorus cocincinensis Duméril and Bibron.)

Aspris BiytH, Journ. Asiatic Soc. Bengal, vol. 22, p. 650, 1853. (Type: Aspris
berdmorei Blyth.)

Enoplosaurus SAuvacE, Bull. Soc. Philom., ser. 7, vol. 3, p. 211, 1879. (Type:
Enoplosaurus insignis Sauvage = Tropidophorus grayt Giinther. )

Eyelids well developed, movable, the lower one scaly and lacking a
more or less transparent or translucent disc ; no supranasals ; prefron-
tals well developed, usually forming a median suture; frontoparietal
single or paired but always distinct from the interparietal ; four prin-
cipal supraoculars; ear large, tympanum superficial; two or three
enlarged preanals ; limbs well developed, digits 5-5, claws nonretractile.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Distribution—Southern China, Indo-China, Philippine Islands,
Australia.
Genus NORBEA Gray
Norbea Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, p. tor, 1845. (Type: Norbea brookei Gray.)
Amphixestus Peters, Monatsb. Akad. Wiss. Berlin, 1871, p. 573. (Type:
Amphixestus beccarii Peters.)

Differs essentially from Tropidophorus as follows: Five principal
supraoculars ; a single greatly enlarged preanal plate.
Distribution —Sarawak, Borneo, Philippine Islands.

Genus TRIBOLONOTUS Duméril and Bibron

Tribolonotus DumérIL and Brsron, Erpétologie générale ..., vol. 5, p. 364,
1839. (Type: Zonurus novae-guineae Schlegel.)

Pediporus Roux, Verh. Naturf. Ges. Basel, vol. 41, p. 129, 1930. (Type:
Tribolonotus schmidti Burt.)

Eyelids well developed, movable, the lower one scaly and lacking
a more or less transparent or translucent disc; head with a posteriorly
emarginate bony casque; nostril in a single nasal; no supranasals ; no
prefrontals ; frontoparietals paired, distinct from the interparietal ; ear
opening approximating eye in size; a pair of greatly enlarged pre-
anals; dorsal surfaces of body and tail armed with large spinose
scales ; neck prominently narrower than the head; limbs large, digits
5-5, claws nonretractile.

Distribution—New Guinea and Solomon Islands.

ALPHABETICAL GENERIC SYNONYMY

Appended below is an alphabetical listing of the 33 generic names
considered valid in this paper, as well as the 62 primary synonyms
of these names which have been proposed over the years. Earlier
names unavailable because of homonymy are listed simply as straight
synonyms of their respective genera. Nomina nuda and new generic
names proposed herein are so indicated; nomina inquirendae are
omitted.

Ablepharus Fitzinger, 1823. Valid genus.

Amphixestus Peters, 1871 = Norbea Gray.

Anomalopus Dumeéril, 1851 = Saiphos Gray.

Anotis Bavay, 1869. Valid genus.

Apterygodon Ederling, 1863 = Dasia Gray.

Aspris Blyth, 1853 = Tropidophorus Duméril and Bibron.
Ateuchosaurus Gray, 1845. Valid genus.

Blepharosteres Stoliczka, 1872—= Ablepharus Fitzinger.
Brachystopus Duméril and Bibron, 1839 = Rhodona Gray.

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 17

Campsodactylus Duméril, 1837 = nomen nudem.
Campsodactylus Duméril and Bibron, 1839 = Riopa Gray.
Carlia Gray, 1844. Valid genus.

Chelomeles Duméril and Bibron, 1839 = Hemiergis Wagler.
Chiamela Gray, 1839 = Riopa Gray.

Coloscincus Peters, 1876 = Saiphos Gray.

Cophoscincopus Mertens, 1934. Valid genus.

Cophoscmcus Peters, 1867. Valid genus.

Cryptoblepharus Wiegmann, 1834. Valid genus.

Cyclodina Girard, 1857 = Leiolopisma Duméril and Bibron.
Dasia Gray, 1839. Valid genus.

Elania Gray, 1845 = Sphenomorphus Fitzinger.

Emoia Gray, 1845. Valid genus.

Enoplosaurus Sauvage, 1879 = Tropidophorus Duméril and Bibron.
Eugongylus Fitzinger, 1843. Valid genus

Eulamprus Fitzinger, 1843 = Sphenomorphus Fitzinger.
Eulepis Fitzinger, 1843 = Lampropholis Fitzinger.
Eumecia Bocage, 1870. Valid genus.

Eusoma Fitzinger, 1843 = Emoia Gray.

Hagria Gray, 1839 = Riopa Gray.

Hemiergis Wagler, 1830. Valid genus.

Heteropus Duméril and Bibron, 1839 = Carlia Gray.
Hinulia Gray, 1845 = Sphenomorphus Fitzinger.

Hombroma Girard, 1857 = Leiolopisma Duméril and Bibron.
Ictiscincus Smith, 1937 = Sphenomorphus Fitzinger.
Isopachys Lonnberg, 1916 = Ophioscincus Peters.
Keneuxia Gray, 1845 = Dasia Gray.

Lamprolepis Fitzinger, 1843 = Dasia Gray.

Lampropholis Fitzinger, 1843. Valid genus.

Leiolopisma Duméril and Bibron, 1839. Valid genus.
Lepidothyris Cope, 1892 = Mochlus Giinther.

Leptosiaphos Schmidt, 1943. Valid genus.

Leptosoma Fitzinger, 1843 = Nodorha, new genus.

Lerista Bell, 1833 = Ablepharus Fitzinger.

Lioscincus Bocage, 1873 = Lampropholis Fitzinger.
Liosoma Fitzinger, 1843 = Tachygyia, new genus.
Liotropis Fitzinger, 1843 = Dasia Gray.

Lipinia Gray, 1845. Valid genus.

Lissonota Blyth, 1853 = Sphenomorphus Fitzinger.
Lygisaurus de Vis, 1884 = Carlia Gray.

Lygosaurus Hallowell, 1860 = Ateuchosaurus Gray.
Lygosoma Hardwicke and Gray, 1827. Valid genus.
Lygosomella Girard, 1857 = Leiolopisma Duméril and Bibron.
Menetia Gray, 1844 = Ablepharus Fitzinger.
Microblepharus Fitzinger, 1843 = Ablepharus Fitzinger.
Miculia Gray, 1844 = Ablepharus Fitzinger.

Mochlus Giinther, 1864. Valid genus.

Mocoa Gray, 1845 = Lampropholis Fitzinger.

Morethia Gray, 1844 = Cryptoblepharus Wiegmann.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS

Myophila de Vis, 1884 = Carlia Gray.
Nannoscincus Ginther, 1872 = Anotis Bavay.
Nodorha, new genus.

Norbea Gray, 1845. Valid genus.

Obligosoma Girard, 1857 = Leiolopisma Duméril and Bibron.

Ophiopsis Fitzinger, 1843 = Ablepharus Fitzinger.
Ophioscincus Peters, 1873. Valid genus.
Otosaurus Gray, 1845. Valid genus.

Panaspis Cope, 1868. Valid genus.

Parotosaurus Boulenger, 1914 = Otosaurus Gray.

Pediporus Roux, 1930 = Tribolonotus Duméril and Bibron.

Peromeles Wiegmann, 1834 = Saiphos Gray.
Peromelis Wagler, 1830 = Hemtergis Wagler.
Phaneropus Fischer, 1881 = Ablepharus Fitzinger.
Pholeophilus Smith, 1849 = Rhodona Gray.

Podophis Wiegmann, 1834 = Lygosoma Hardwicke and Gray.

Praepeditus Duméril and Bibron, 1839 = Rhodona Gray.
Rhodona Gray, 1839. Valid genus.

Riopa Gray, 1839. Valid genus.

Ristella Gray, 1839. Valid genus.

Ronia Gray, 1841 = Rhodona Gray.

Saiphos Gray, 1831. Valid genus.

Sauroscincus Peters, 1879 = Lampropholis Fitzinger.
Scincella Mittleman, 1950. Valid genus.

Sepacontias Giinther, 1880 = Mochlus Giinther.
Soridia Gray, 1839 = Rhodona Gray.

Sphenomorphus Fitzinger, 1843. Valid genus.
Sphenosoma Fitzinger, 1843 = Riopa Gray.
Squamicilia, new genus.

Tachygyia, new genus.

Tetradactylus Cuvier, 1829 = Hemiergis Wagler.
Theconyx Annandale, 1906 = Dasia Gray.

Tribolonotus Dumeéril and Bibron, 1839. Valid genus.
Tridactylus Cuvier, 1829 = Hemiergis Wagler.
Tropidophorus Duméril and Bibron, 1839. Valid genus.
Tropidoscincus Bocage, 1873 = Lampropholis Fitzinger.
Typhloseps Angel, 1920 = Ophioscincus Peters.

KEY TO THE GENERA OF LIZARDS OF THE SUBFAMILY

LYGOSOMINAE

1. Head broad and distinct from the relatively slender neck, with
a posteriorly emarginate bony casque; dorsum of body and

: £07

tail with large spinose scales.......... Tribolonotus Duméril and Bibron

Head not prominently distinct from neck ; no bony casque; body

and tail not covered with large spinose scales..................
2. Byehdsvabsenteand/orm 1mmoyablese nite eee eee eee ene
Eyelids present and)movablevr rcs acire ic cis bse teisiaeies cyeiniee eee

3. No upper eyelid, lower lid immovable and without a more or

less transparent or translucent disc; no limbs...... Ophioscincus Peters

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 19

10.

TY.

12.

13;

14.

15.

16.

D7.

Upper eyelid present but often much reduced and more or less

fused with the lower lid which bears a large transparent or

Lranisinceme aise’, HMDS resenby/ 60.5. Soule cael beck Rete ae 4
Frontoparietal single or paired, but always distinct from the

interparietal; ear opening absent, or smaller than the eye if

PUCSEHED sie Cyoem amt ati a aia wore OS.o See a awe ee ce ae Ablepharus Fitzinger
Frontoparietal and interparietal fused to form a single large

shield ; ear opening approximates eye opening. . Cryptoblepharus Wiegmann
Lower eyelid scaly, lacking a more or less transparent or

translucent disc; occasionally with a small opaque disc............. 6
Lower eyelid with a more or less transparent or translucent disc..... 21
Supranasals present (sometimes fused partially with the nasals)...... 7
SAPEANASALS SCO rc Natalee. Pars atcrale «e ‘aboriliure abil laiepiany aie mete ace aT eae 13
Baragening: presents vo. <0 Aas. os Cee ee, Cee eee 8
Ear opening absent, or else covered by overlapping scales............ 12
Preanals prominently enlarged; ear opening approximates the

GIAMELERH OP CLOT EY OOF ewe u ete iana eRe diate ne Ga oe Otosaurus Gray
Preanals not, or barely, enlarged; diameter of ear opening half

orlesshinediametenr'of ‘the eye. s isan. 606i. Se. oceoe dene Oe aca 9
Supranasals reduced, separated by the rostro-frontonasal suture........ 10
Supranasals enlarged, forming a median suture...................00. II

Frontoparietals paired; lower eyelid usually uniformly scaly

but occasionally with a small opaque disc; 5 supraoculars;

limbs erobustaOdy StOuUtcreiac aciteeiys nielele ste ciciterel Eugongylus Fitzinger
Frontoparietals united; lower eyelid uniformly scaly, never

with a small opaque disc; 4 supraoculars; limbs much re-

duced and weak, body slender and vermiform....Squamicilia, new genus
Frontoparietals paired; limbs short, barely meeting or sepa-

rated by several scale lengths when appressed........ Mochlus Gunther
Frontoparietals united; limbs long, widely overlapping when

APBLESSCE Ae sists cg cocastaec sd Maks recite wae salve Tachygyia, new genus
Preanals not, or barely, enlarged; supranasals not, or rarely,

fOrmin ed Median SULUTes. 2.6... 05 25. 2 ead. chew aeleh cane Dasia Gray
Preanals prominently enlarged ; supranasals large and forming a

Hedian SUeUTe Ses ahs Ge ae Pea eae eee ewe Cophoscincopus Mertens

True parietals lacking; frontal very long and constricted,
longer than frontoparietals plus interparietal; prefrontals

SHIANSEDATALE Sistyls sles valerie s Sass ee cimenemobn Ateuchosaurus Gray
True parietals present ; frontal not long and constricted, shorter

than frontoparietals plus interparietal. /.....:...000ccesbeebaea ce. 14
Ryman SUPECNCIAl: J... oases et elec aeeicy swe eoRwineeecaesbucmem ay 15
Tympanum sunk (if visible at all). ........... cece cen weeecneeecrene 16

Enlarged preanals 2 or 3; supraoculars 4.
Tropidophorus Duméril and Bibron

Enlarged preanal £3 supraoculars 5... 6.0... ees cn ee nneaes Norbea Gray
PrOntaparictals WME oc. eck oleae sa cedew scenes te egeenessnelngs 17
rontoparietals) PAITEM <....eccc csv uieeeoccedo es awit ew rinseeis epmsie ee 18

Preanals not, or barely, enlarged; ear opening usually present

but punctiform; limbs much reduced....Lygosoma Hardwicke and Gray
Preanals enlarged; ear opening absent; limbs robust, often

meeting or overlapping when appressed...........- Cophoscincus Peters

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

28.

20.

30.

31.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 17

Claws: retrambile 1a jisbaiags Hawes bee orou Haybbre hea ence nens Ristella Gray

Claws: nonretractile «nk ccuuds shawians saleaty a ewakiet: Sake eek ee 19
Prefrontals large, often forming a median suture; limbs robust;

5 Wi gttel)--aecena te oteraces sattas ctteamtees eee ncearsia cian biavs Sphenomorphus Fitzinger
Prefrontals greatly reduced and widely separated, or else

absent; digits 5 or less, limbs reduced and weak...............05 20
Preanals enlarged; supraoculars 3 or 4; ear opening absent;

Cae Sah atta gues N oy Pike DORN te, hil; Regt Niel MWe ak rue ate Saiphos Gray

Preanals not, or barely, enlarged; supraoculars 4; ear opening
present although often much reduced; digits 3, 4, or 5.
Leptosiaphos Schmidt

Supranasals present (sometimes fused partially with the nasals)...... 22
Supranasals absents sua, semua ecroetieciale 6 eile ee aD ae eee eee 25
Supranasals large, forming a median suture................ cece eee 23
Supranasals reduced, separated by the rostro-frontonasal suture........ 24
Prefrontals large, usually forming a median suture or else

barely separated by the rostro-frontonasal suture; ear ap-

proximates eye in size; limbs vestigial, digits 2 or 3....Ewmecia Bocage
Prefrontals small, widely separated; ear opening approximates

size of nostril, or smaller; limbs short but robust, digits 3 or

TTLOT G25 tallot Sra catia ionatter tee tena atecSUalstols cous tuare vw rGiehe etonere eas tetcns: siGnetere Riopa Gray
Frontoparietals united and usually fused with the interparietal

to form a single large shield, or else if the interparietal is

present it is much reduced; limbs large and broadly over-

lapping when appressed; ear approximates nostril in size..Emoia Gray
Frontoparietals paired and distinct from the well-developed

interparietal; limbs separated, just meeting, or overlapping

when appressed; ear large, approximating eye in size....Panaspis Cope

Preanals snot; (or ‘barely; enlarged. ich, < ld acehocia vic sree = bse hopetaye or ciouemaioial 26
Preanaissprominentlyenlargedicisi:. asictecteteyeinte)< dates ceyerosels jee hay 29
Frontoparietalss Waited cic5.c:.ccssyaceyevaieteieseyoceusl sissies 0 obs mal s)aysueiv eaierevaraaiato oh 27
Frontopatictals «paired, 2ictiasaihnss <n balemets careers nye, sts e)a krone a 28
Interparietal distinct, usually large; digits 5-5....Lampropholis Fitzinger
Interparietal very small or absent; digits 4-5..............05- Carlia Gray

Digits 5-5; limbs well developed, meeting or overlapping when

appressed, or else separated by one or two scale lengths; ear

OPEN Ne HPrOMIUINENE. flac, Leilaietaisterare oe Mee evel Leiolopisma Duméril and Bibron
Digits 5-5 or less; limbs short, separated by several scale

lengths when appressed; ear opening absent or punctiform

(approximating ‘nostril or smaller) ..o.0 4 sm -islecin erento Anotis Bavay
Frontoparietals and interparietal fused to form a single large
shield; anterior limbs always lacking.................+- Rhodona Gray

Frontoparietals united or paired, but always distinct from
interparietal; anterior limbs present (except in Nodorha

EMERGE) A wicikie ce hiteele otiare wie hee seine pace ete rece e sue pat Orage Cae ale Var tn ea 30
Limbs much reduced or rudimentary, digits 5-5 or less..............6. 31
Limbs well developed, often meeting or overlapping when ap-

PECSSE s ISLES ALWAYS \525 oa. wie alee. ajeteinia je aie Giale wo wlainh tn seine eres 32

Prefrontals large, often forming a median suture; nasals small
to moderate in size, usually separated.............+- Hemiergis Wagler

NO. I7 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 21

Prefrontals small and widely separated, or absent; nasals large,

usually forming a median suture................6. Nodorha, new genus
ae BAG tCOUCCAIEd! etciisisles & tlecrsteS oieleeMe dine asmtreslaieales Cophoscincus Peters
Babe ODeHing 1 PeeSeBty < jase, nhs aie ual w sorte hens s capaldye bore eta sea 33

33. Prefrontals forming a median suture longer than half their

length; frontonasal at least as long as it is wide, or longer ;

snout elongated, acute, depressed or concave; rostral deeper
(Higher) thant) 19“ wides.(.s 22. kkids «ciwiams slnntoste oan aes Lipinia Gray

Prefrontals separated or forming a median suture less than half

their length; frontonasal wider than it is long; snout short,
obtuse, flat or convex; rostral wider than it is high..Scincella Mittleman

ALPHABETICAL LIST OF SPECIES

In the listing that follows, the fundamental intent is to indicate the
limits of the genera employed in this paper, through the medium of
their included species. In no sense does this list purport to be a check-
list of the forms considered valid, nor does it enumerate all the names
that have been proposed in the Lygosominae. Considerable latitude
regarding the inclusion of a name has been exercised; where a form
has been rather universally regarded as a synonym of an older name,
the general rule has been to omit the junior name. On the other hand,
some names have been included despite their relegation to synonymy
by other workers, since in these cases it is believed likely that further
study will demonstrate a distinguishable racial entity.

acrocarinata Kopstein, Emota, 1926.

acutus Peters, Sphenomorphus, 1864.
adelaidensis Peters, Cryptoblepharus, 1874.
adspersa Steindachner, Emoia, 1870.

aenea Girard, Leiolopisma, 1857.

aeneus Cope, Panaspis, 1868.

aerata Garman, Carlia, 1901.

aestuosa Girard, Leiolopisma, 1857.
africana Gray, Leiolopisma, 1845.
africanus Sternfeld, Cryptoblepharus, 1918.
ahli Vogt, Emoia, 1932.

aignanus Boulenger, Sphenomorphus, 1808.
albertisii Peters and Doria, Carlia, 1878.
albofasciolatus Giinther, Eugongylus, 1872.
albopunctatus Gray, Mochlus, 1846.
aldabrae Sternfeld, Cryptoblepharus, 1918.
alfredi Boulenger, Sphenomorphus, 1808.
aloysii-sabaudiae Peracca, Leptostaphos, 1907.
amblyplacodes Vogt, Sphenomorphus, 1932.
anchietae Bocage, Ewmecia, 1870.

anguina Theobald, Riopa, 1868.
anguinoides Boulenger, Ophioscincus, 1914.
annamiticus Boettger, Sphenomorphus, 1901.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

annectens Boulenger, Otosaurus, 1897.

anolis Boulenger, Lipinia, 1883.

anomalopus Boulenger, Sphenomorphus, 1800.
anomalus Gray, Cryptoblepharus, 1844.
antoniorum Smith, Sphenomorphus, 1927.
aruanus Roux, Sphenomorphus, 1910.
aruensis Doria, Sphenomorphus, 1874.
aruensis Sternfeld, Emoia, 1918.

assamensis Annandale, Tropidophorus, 1912.
assata Cope, Scincella, 1864.

ater Boettger, Cryptoblepharus, 1913.
atrigulare Ogilby, Carlia, 1890.

atrigularis Stejneger, Sphenomorphus, 1908.
atrocostata Lesson, Emoia, 1830.
atromaculatus Garman, Sphenomorphus, 1901.
aubrianus Bocage, Lampropholis, 1873.
australe Gray, Lygosoma, 1839.

australe Peters, Ophioscincus, 1873.

australis Gray, Sphenomorphus, 1839.
autralis Sternfeld, Cryptoblepharus, 1918.
austro-caledonicus Bavay, Lampropholis, 1860.
balinensis Barbour, Cryptoblepharus, 191t.
bampfylede: Bartlett, Mochlus, 1895.
bancrofti Longman, Saiphos, 1916.

barbouri Stejneger, Scincella, 1925.

battersbyt Procter, Emoia, 1920.

baudini Duméril and Bibron, Emoia, 1830.
beauforti de Jong, Sphenomorphus, 1927.
beccarti Peters, Norbea, 1871.

beccarit Peters and Doria, Carlia, 1878.
beddomii Boulenger, Ristella, 1887.

beddomii Boulenger, Scincella, 1887.
berdmorei Blyth, Tropidophorus, 1853.
bicarinata Macleay, Carlia, 1877.

bilineata Gray, Scincella, 1846.

biparietalis Taylor, Sphenomorphus, 1918.
bipes Fischer, Rhodona, 1882.

bitaeniatus Boettger, Cryptoblepharus, 1913.
biunguiculatus Oudemans, Saiphos, 1894.
bivittatus Menestries, Ablepharus, 1832.
blackmanni de Vis, Carlia, 1885.

blanchardi Burt, Tribolonotus, 1930.
blochmanni Tornier, Leptosiaphos, 1903.
boetigeri Van Denburgh, Scincella, 1912.
boettgeri Sternfeld, Emoia, 1918.

boettgert Sternfeld, Eugongylus, 1918.
bougainvillii Duméril and Bibron, Nodorha, 1839.
boulengert Van Denburgh, Sphenomorphus, 1912.
bouton Desjardins, Cryptoblepharus, 1831.
bowringi Ginther, Mochlus, 1864.

117

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN

brachysomus Lonnberg and Andersson, Sphenomorphus, 1915.
brandti Strauch, Ablepharus, 1868.

breviceps Peters, Panaspis, 1873.

brevipes Boettger, Sphenomorphus, 1895.
brookei Gray, Norbea, 1845.

brooksi Loveridge, Sphenomorphus, 1933.
browni Van Denburgh, Ateuchosaurus, 1912.
buergersi Vogt, Emota, 1932.

burdeni Dunn, Cryptoblepharus, 1927.
burgeoni Witte, Leptosiaphos, 1933.
buttikoferi Lidth de Jeude, Sphenomorphus, 1905.
cabindae Bocage, Panaspis, 1866.
cacrulecauda de Vis, Emoia, 1802.

callisticta Peters and Doria, Emoia, 1878.
cameronicus Smith, Sphenomorphus, 1924.
caudaequinae Smith, Scincella, 1950.

caudatus Sternfeld, Cryptoblepharus, 1918.
celebense Miiller, Otosaurus, 1804.

celebense de Rooij, Dasia, 1915.

challengeri Boulenger, Leiolopisma, 1887.
cherriei Cope, Scincella, 1893.

chinensis Gray, Ateuchosaurus, 1845.
cocincinensis Duméril and Bibron, Tropidophorus, 1839.
cognatus Boettger, Cryptoblepharus, 1881.
colletti Boulenger, Sphenomorphus, 1896.
compressicauda Witte, Leptosiaphos, 1933.
comtus Roux, Sphenomorphus, 1928.
concinnatus Boulenger, Otosaurus, 1887.
consobrinus Peters and Doria, Sphenomorphus, 1878.
cophia Boulenger, Lygosoma, 1908.
corpulentus Smith, Mochlus, 1921.

courcyanus Annandale, Sphenomorphus, 1912.
crassicaudus Duméril, Sphenomorphus, 1851.
cumingi Gray, Otosaurus, 1845.

cuneiceps de Vis, Emoia, 1890.

cuprea Gray, Leiolopisma, 1839.

cursor Barbour, Cryptoblepharus, 1911.

curta Boulenger, Carlia, 1897.

curtirostris Taylor, Otosaurus, 1915.
cyanogaster Lesson, Emoia, 1830.

cyanura Lesson, Emoia, 1830.

dahomeyense Chabanaud, Panaspis, 1917.
darlingtoni Loveridge, Lygosoma, 1933.
darlingtoni Loveridge, Tropidophorus, 1945.
decipiens Boulenger, S. phenomorphus, 1894.
decresiensis Fitzinger, H emiergis, 1829.
degrijsi Mertens, Cryptoblepharus, 1928.
delicata de Vis, Lampropholis, 1888.

dendyi Boulenger, Leiolopisma, 1902.
deplanchti Bavay, S ‘phenomorphus, 1869.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

derooyae de Jong, Sphenomorphus, 1927.
deserti Strauch, Ablepharus, 1868.

desertora Sternfeld, Nodorha, 19109.

devisti Boulenger, Carlia, 1890.

diguliense Kopstein, Carlia, 1926.

distinguendus Werner, Ablepharus, 1910.
divergens, Taylor, Sphenomorphus, 1920.
domina de Vis, Sphenomorphus, 1888.

doriae Boulenger, Scincella, 1887.

dorsalis, Boulenger, Sphenomorphus, 1887.
durus Cope, Cophoscincopus, 1862.

dussumieri Duméril and Bibron, Sphenomorphus, 1839.
elberti Sternfeld, Dasia, 1918.

elegans Gray, Ablepharus, 1844.

elegans Boulenger, Scincella, 1897.

elegans Sternfeld, Sphenomorphus, 1918.
elegantoides Ahl, Scincella, 1925.

elegantulus Peters and Doria, Sphenomorphus, 1878.
emigrans Lidth de Jeude, Sphenomorphus, 1895.
entrecasteauxit Duméril and Bibron, Leiolopisma, 1830.
equalis Gray, Saiphos, 1825.

essingtont Gray, Sphenomorphus, 1842.
euryotts Werner, Lampropholis, 1909.

exigua Anderson, Scincella, 1878.

fallax Peters, Sphenomorphus, 1860.

fasciatus Gray, Sphenomorphus, 1845.

fasciolara Girard, Leiolopisma, 1857.

fasciolatus Giinther, Sphenomorphus, 1867.
fernandt Burton, Mochlus, 1836.

fischeri Boulenger, Sphenomorphus, 1887.
flavigulare Schmidt, Emoia, 1932.

flavipes Parker, Sphenomorphus, 1936.

florense Weber, Sphenomorphus, 1891.

foliora de Vis, Carlia, 1884.

forbest Boulenger, Sphenomorphus, 1888.
forbesora Taylor, Scincella, 1937.

formosa Blyth, Leiolopisma, 1853.

formosensis Van Denburgh, Scincella, 1912.
formosensis Van Denburgh, Sphenomorphus, 1912.
fragile Giinther, Nodorha, 1876.

frontalis de Vis, Ophioscincus, 1888.

frosti Zeitz, Nodorha, 1920.

furcatus Weber, Cryptoblepharus, 1801.

fusca Dumeéril and Bibron, Carlia, 18309.

garniert Bavay, Tachygyia, 18609.

gemmingeri Cope, Scincella, 1864.

gerrardii Gray, Nodorha, 1864.

gloriosus Stejneger, Cryptoblepharus, 18093.
gracile Bavay, Anotis, 1860.

gracilis de Rooij, Tribolonotus, 1909.

117

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN

graciloides Lonnberg and Andersson, Anotis, 1913.
grande Gray, Leiolopisma, 1845.

granulatus Boulenger, Otosaurus, 1903.
graueri Sternfeld, Leptosiaphos, 1912.
grayanus Stoliczka, Ablepharus, 1872.

grayi Gtinther, Tropidophorus, 1861.

greyit Gray, Ablepharus, 1844.

grifint Taylor, Dasia, 1915.

guentheri Peters, Riopa, 1879.

guentheri Boulenger, Ristella, 1887.
guichenoti Duméril and Bibron, Lampropholis, 1839.
guineensis Peters, Mochlus, 1879.
gyldenstolpei Lonnberg, Ophioscincus, 1916.
hainanus Smith, Tropidophorus, 1923.
haliana Haly and Nevill, Dasia, 1887.
hallieri Lidth de Jeude, Sphenomorphus, 1905.
helenae Cochran, Sphenomorphus, 1927.
helleri Loveridge, Leptosiaphos, 1932.
herberti Smith, Mochlus, 1916.

himalayana Giinther, Scincella, 1864.
incerta Stuart, Scincella, 1940.

inconspicua Miiller, Scincella, 18094.

indicus Gray, Sphenomorphus, 1853.
infralineolatus Giinther, Cophoscincus, 1873.
infrapunctata Boulenger, Leiolopisma, 1887.
iniqua Lidth de Jeude, Norbea, 1905.

initialis Werner, Anotis, 1910.

inornatus Gray, Sphenomorphus, 1845.
intermedius Sternfeld, Sphenomorphus, 1919.
intermedius Kinghorn, Sphenomorphus, 1932.
iridescens Boulenger, Emoia, 1897.

irrorata Macleay, Emoia, 1877.

isodactyla Giinther, Squamicilia, 1864.
isolepis Boulenger, Sphenomorphus, 1887.
ixbaac Stuart, Scincella, 1940.

jagori Peters, Sphenomorphus, 1864.

jakati Kopstein, Emoia, 1926.

jamnana Loveridge, Carlia, 1948.

jeudei Boulenger, Sphenomorphus, 1807.
jobiensis Meyer, Sphenomorphus, 1874.
johnstoni Boulenger, Ewmecia, 1897.
keiensis Roux, Cryptoblepharus, 1910.
keiensis Sternfeld, Emoia, 1918.

kilimensis Stejneger, Leptosiaphos, 180t.
kinabaluensis Bartlett, Otosawrus, 1895.
kitsont Boulenger, Panaspis, 1913.

Rlossi Boulenger, Emoia, 1914 .

kohtaoensis Cochran, Scincella, 1927.
koratensis Smith, Mochlus, 1917.

kordoana Meyer, Emoia, 1874.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

kosciuskoi Kinghorn, Sphenomorphus, 1932.
kuekenthali Boettger, Emoia, 1895.

kuhneit Roux, Sphenomorphus, 1910.

kutuensis Lonnberg, Leptosiaphos, 1911.
labillardierii Gray, Sphenomorphus, 1838.
ladacense Giinther, Scincella, 1864.

laeve Oudemans, Carlia, 18904.

laeviceps Peters, Mochlus, 1874.

laotus Smith, Tropidophorus, 1923.

larutense Boulenger, Saiphos, 1900.

laterale Say, Scincella, 1823.

lateralis de Vis, Carlia, 1885.

laterimaculata Boulenger, Scincella, 1887.
latifasciatus Meyer, Sphenomorphus, 1874.
leae Boulenger, Sphenomorphus, 1887.
lentiginosus de Vis, Saiphos, 1888.

leonhardii Sternfeld, Sphenomorphus, 1910.
leschenaultu Cocteau, Cryptoblepharus, 1832.
lesweuru. Duméril and Bibron, Sphenomorphus, 1839.
leucospilus Peters, Tropidophorus, 1872.
leucotaema Bleecker, Carlia, 1860.

leveretti Schmidt, Sphenomorphus, 1927.
lichenigera O’Shaughnessy, Leiolopisma, 1874.
lineata Gray, Rhodona, 1839.

lineata Gray, Riopa, 1830.

lineatus Bell, Ablepharus, 1833.

lineolata Stoliczka, Riopa, 1870.

lineo-ocellata Duméril, Leiolopisma, 1851.
lineo-ocellatus Duméril and Bibron, Cryptoblepharus, 1839.
lineopunctulata Duméril and Bibron, Rhodona, 1839.
llanosi Taylor, Otosaurus, 1910.

lobula Loveridge, Scincella, 1945.
longicaudatus de Rooij, Sphenomorphits, 1915.
longiceps Boulenger, Lipinia, 1895.

loriae Boulenger, Sphenomorphus, 1897.
louisadensis Boulenger, Sphenomorphus, 1003.
luberoensis Witte, Leptosiaphos, 1933.
luctuosa Peters and Doria, Carlia, 1878.
luzonense Boulenger, Lygosoma, 1895.
mabuiforma Loveridge, Riopa, 1935.
maccooeyi Ramsay and Ogilby, Carlia, 1890.
maccoyi Lucas and Frost, Anotis, 1894.
macropisthopa Werner, Nodorha, 1903.
macrota Steindachner, Scincella, 1869.
macrotympana Stoliczka, Scincella, 1873.
maculatus Blyth, Sphenomorphus, 1853.
maindront Sauvage, Sphenomorphus, 1879.
malayanus Doria, Sphenomorphus, 1888.
manm Brown, Emoia, 1948.

mariae Bavay, Anotis, 1860.

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN

mayottensis Mertens, Cryptoblepharus, 1928.
megalops Annandale, Sphenomorphus, 1906.
megaspilus Gunther, Sphenomorphus, 1877.
mehelyt Werner, Emoia, 1899.

melanochlorus Vogt, Sphenomorphus, 1932.
melanopogon Dumeéril and Bibron, Sphenomorphus, 1830.
melanopogon Gray, Carlia, 1844.

melanosticta Boulenger, Scincella, 1887.
meleagris Boulenger, Leptosiaphos, 1907.
mentovarius Boettger, Eugongylus, 1895.
metallicum O’Shaughnessy, Lampropholis, 1874.
metallicus Boulenger, Cryptoblepharus, 1887.
miangense Werner, Lipinia, 1910.

microcerca Boettger, Scincella, 1901.
microlepidota O'Shaughnessy, Lampropholis, 1874.
microlepis Duméril and Bibron, Tachygyia, 1839.
microlepis Giinther, Tropidophorus, 1861.
micropa Lidth de Jeude, Norbea, 10905.
microta Gray, Nodorha, 1844.

mimikanus Boulenger, Otosaurus, 1914.
minutus Meyer, Sphenomorphus, 1874.
miodactylus Boulenger, Saiphos, 1903.

miopa Ginther, Rhodona, 1867.

miota Boulenger, Scincella, 1805.

misaminia Stejneger, Norbea, 1908.

mivarti Boulenger, Emoia, 1887.

myjobergt Lonnberg, Sphenomorphus, 1913.
moco Duméril and Bibron, Letolopisma, 18309.
mocquardi Boulenger, Norbea, 1894.
mocquardi Chabanaud, Mochlus, 1918.
modesta Gunther, Scincella, 1864.

modestus Gunther, Mochlus, 1880.

modigliant Boulenger, Sphenomorphus, 1805.
moellendorffii Boettger, Sphenomorphus, 1897.
moluccara Barbour, Dasia, Iott.

monotropis Boulenger, Sphenomorphus, 1887.
monticola Schmidt, Scincella, 1927.

morokana Parker, Scincella, 1936.
moszkowskti Vogt, Sphenomorphus, 1912.
moultonit Barbour and Noble, Dasia, 1912.
muelleri Schlegel, Sphenomorphus, 1837.
muelleri Fischer, Ablepharus, 1881.

munda de Vis, Carlia, 1885.

mundivense Browne, Carlia, 1808.

murphyt Burt, Emoia, 1930.

murrayi Boulenger, Sphenomorphus, 1887.
murudensis Smith, Otosaurus, 1925.

mustelina O’Shaughnessy, Leiolopisma, 1874.
neuhaussi Vogt, Sphenomorphus, 191l.
nichollsi Loveridge, Nodorha, 1933.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

nieuwenhuisi Lidth de Jeude, Dasia, 1905.
nigra Hombron and Jacquinot, Emoia, 1853.
nigra Sternfeld, Dasia, 1918.

nigricaudis Macleay, Sphenomorphus, 1877.
nigrigulare Boulenger, Carlia, 1897.
nigriventris de Rooij, Sphenomorphus, 1915.
migrofasciolata Peters, Letolopisma, 1869.
nigrolabrus Giinther, Sphenomorphus, 1873.
nigrolineatus Boulenger, Sphenomorphus, 1897.
nigropunctatus Hallowell, Cryptoblepharus, 1860.
nimbense Angel, Panaspis, 1944.

nitens Peters, Scincella, 1871.

noctua Lesson, Lipinia, 1830.

nototaenius Boulenger, Sphenomorphus, 1914.
novaecaledomiae Parker, Lampropholis, 1926.
novaeguineae Schlegel, Tribolonotus, 1834.
novaeguineae Meyer, Carlia, 1874.
novaeguineae Mertens, Cryptoblepharus, 1928.
novocaledonicus Mertens, Cryptoblepharus, 1928.
obscura de Jong, Emoia, 1927.

ocellata Gray, Lampropholis, 1844.

ocellatus Boulenger, Sphenomorphus, 1806.
ocelliferus Boulenger, Sphenomorphus, 1806.
oligolepis Boulenger, Sphenomorphus, 1914.
olivacea Gray, Dasia, 1838.

opisthorhodus Werner, Mochlus, 1910.
orientale Shreve, Sphenomorphus, 1940.
ornatus Gray, Sphenomorphus, 1845.
pagenstecheri Lindholm, Scincella, 1901.
pallidiceps de Vis, Emoia, 1890.

pallidus Gunther, Sphenomorphus, 1844.
pallidus Mertens, Cryptoblepharus, 1928.
palnica Boettger, Scincella, 18092.

pannonicus Fitzinger, Ablepharus, 1823.
papuensis Macleay, Sphenomorphus, 1877.
paraenea Ahl, Leiolopisma, 1925.

pardalis Macleay, Saiphos, 1877.

parietale Peters, Emoia, 1871.

parkeri Smith, Scincella, 1937.

partellot Stejneger, Norbea, 1910.

parvus Boulenger, Sphenomorphus, 18097.
pectorale de Vis, Carlia, 1885.

pellopleurus Hallowell, Ateuchosaurus, 1860.
pembanus Boettger, Mochlus, 1913.

peronii Fitzinger, Hemiergis, 1826.

peronit Cocteau, Cryptoblepharus, 1836.
perontti Duméril and Bibron, Carlia, 1839.
perplexa Barbour, Norbea, 1921.

perviridis Barbour, Dasia, 1921.

phaeodes Vogt, Scincella, 1932.

117

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN

philippinica Mertens, Dasia, 1920.

physicae Duméril and Bibron, Emoia, 1839.
picturata Fry, Nodorha, 1914.

plagiocephalus Cocteau, Cryptoblepharus, 1836.
planivenirale Lucas and Frost, Nodorha, 1902.
poecilopleurus Wiegmann, Cryptoblepharus, 1835.
popae Shreve, Riopa, 1940.

potanini Ginther, Scincella, 1806.

pranensis Cochran, Scincella, 1930.

pratti Boulenger, Sphenomorphus, 1903.
prehensicauda Loveridge, Scincella, 1945.
presignis Boulenger, Sphenomorphus, 1900.
pretiosum. O’Shaughnessy, Lampropholis, 1874.
producta Boulenger, Squamicilia, 1909.
pseudotropa Giinther, Leiolopisma, 1844.
pulchella Gray, Lipinta, 1845.

pulcher Sternfeld, Cryptoblepharus, 1918.
pulchra Boulenger, Scincella, 1903.

pulla Barbour, Carlia, 1911.

pumila Boulenger, Lygosoma, 1887.

punctata Linné, Riopa, 1766.

punctata Gray, Rhodona, 1839.

punctatolineata Boulenger, Scincella, 1893.
punctatovittata Giinther, Nodorha, 1867.
punctatus Sternfeld, Cryptoblepharus, 1918.
punctulata Peters, Lygosoma, 1871.
quadridigitatus Werner, Hemiergis, 1910.
quadrilineata Duméril and Bibron, Hemiergis, 1839.
quadrivittatus Peters, Cophoscincus, 1867.
quadrupes Linné, Lygosoma, 1766.
quatuordecimlineatus Sternfeld, Sphenomorphus, 1919.
quatuordigitata Sternfeld, Leptosiaphos, 1912.
queenslandiae de Vis, Tropidophorus, 1890.
quinquetaeniatus Giinther, Cryptoblepharus, 1874.
quoyi Duméril and Bibron, Sphenomorphus, 1830.
reevesi Gray, Scincella, 1838.

reichenovei Peters, Leiolopisma, 1874.

relictus Vinciguerre, Cophoscincus, 1892.

renschi Mertens, Cryptoblepharus, 1928.
reticulatus Giinther, Saiphos, 1873.

rhomboidale Peters, Carlia, 1860.

richardsoni Gray, Sphenomorphus, 1844.
rivulare Taylor, Norbea, 1015.

robinsoni Smith, Tropidophorus, 19109.

roulei Angel, Ophioscincus, 1920.

rouxi Hediger, Lipinia, 1934.

rufescens Shaw, Eugongylus, 1802.

ruficauda Taylor, Emoia, 1915.

ruficaudus Lucas and Frost, Cryptoblepharus, 1895.
rufus Boulenger, Sphenomorphus, 1887.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. TE

rupicola Smith, Scincella, 1916.

rurku Gray, Ristella, 1839.

rutilus Peters, Crytoblepharus, 1879.
salsburyi Schmidt, Ateuchosaurus, 1927.
samoensis Duméril, Emoia, 1851.

sanctus Dumeéril and Bibron, Sphenomorphus, 1830.
sanfordi Schmidt and Burt, Emoia, 1930.
sarasinorus Boulenger, Sphenomorphus, 1897.
scharffi Boulenger, Anotis, 1915.

schevilli Loveridge, Sphenomorphus, 1933.
schmidti Burt, Tribolonotus, 10930.

schoedei Vogt, Sphenomorphus, 1912.
schultzei Vogt, Sphenomorphus, 1911.
scotophilus Boulenger, Sphenomorphus, 1900.
scutirostra Peters, Lygosoma, 1873.
sembalunica Mertens, Carlia, 1927.

semoni Oudemans, Scincella, 1804.

semperi Peters, Scincella, 1867.
septentrionale Schmidt, Scincella, 1927.
shelfordi Boulenger, Sphenomorphus, 1900.
sikkimense Blyth, Scincella, 1854.

silvicola Taylor, Scincella, 1937.

similis Dunn, Emota, 1927.

simplex Cope, Saiphos, 1864.

simus Sauvage, Sphenomorphus, 1879.
sinicus Boettger, Tropidophorus, 1886.
slevini Loveridge, Anotis, 1941.

smaragdina Lesson, Dasia, 1830.

smithi Gray, Leiolopisma, 1845.

solomonis Boulenger, Sphenomorphus, 1887.
sorex Boettger, Emoia, 1895.

sowerbyi Stejneger, Ateuchosaurus, 1924.
spaldingi Macleay, Sphenomorphus, 1877.
speisert Roux, Emoia, 1913.

spinaure Smith, Carlia, 1927.

spurrelli, Boulenger, Panaspis, 1917.
stanleyana Boulenger, Scincella, 1897.
steeret Stejneger, Sphenomorphus, 1908.
steindachnert Bocage, Lampropholis, 1873.
siejnegert Taylor, Norbea, 1922.

stellatus Boulenger, Sphenomorphus, 1900.
stickeli Loveridge, Sphenomorphus, 1948.
strauchit Boulenger, Sphenomorphus, 1887.
striatopunctulatus Ahl, Sphenomorphus, 1925.
striolatus Weber, Sphenomorphus, 1891.
stuarti Smith, Scincella, 1941.

subcaerulea Boulenger, Dasia, 1801.
subnitens Boettger, Scincella, 1896.

sulaense Kopstein, Eugongylus, 1927.
sumatrense Bleecker, Saiphos, 1860.

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 31

sumbawanus Mertens, Cryptoblepharus, 1928.
sundevalli Smith, Mochlus, 1849.

surdus Boulenger, Cophoscincus, 1900.

suteri Boulenger, Leiolopisma, 1906.
taentolatus Shaw, Sphenomorphus, 1700.
taemopleurus Peters, Cryptoblepharus, 1874.
tamburinensis Lonnberg and Andersson, Sphenomorphus, 1915.
tanae Loveridge, Riopa, 1935.

taprobanensis Kelaart, Sphenomorphus, 1852.
tavesae Smith, Scincella, 1935.

taylorit Burt, Otosaurus, 1930.

taylort Oliver, Scincella, 1937.

telfairii Desjardins, Letolopisma, 1831.

tenue Garman, Sphenomorphus, 1001.
tenuiculus Mocquard, Sphenomorphus, 1890.
tenuis Gray, Sphenomorphus, 1831.
terdigitata Parker, Nodorha, 1926.

tersus Smith, Sphenomorphus, 1916.
tetradactyla Lucas and Frost, Nodorha, 1875.
tetradactyla O’Shaughnessy, Carlia, 1870.
tetrataenia Boulenger, Emoia, 1895.

texta Mueller, Scincella, 1894.

thai Smith, Tropidophorus, 1910.

thomasi Tornier, Leptosiaphos, 1903.
tigrinus de Vis, Sphenomorphus, 1888.
timidus de Vis, Ablepharus, 1888.

togoensis Werner, Panaspis, 1902.

torniert Vogt, Sphenomorphus, 191.
totocarinatus Vogt, Sphenomorphus, 1932.
travancorica Beddome, Ristella, 1870.
travankorica Beddome, Scincella, 1870.
tricarinata Meyer, Carlia, 1874.

tricolor Bavay, Lampropholis, 1869.
tridactylus Boulenger, Hemiergis, 1915.
trilineatum Gray, Lampropholis, 1838.
tropidolepis Boulenger, Emoia, 1914.
tropidonotus Boulenger, Sphenomorphus, 18097.
truncatus Peters, Saiphos, 1876.

tryoni Longman, Sphenomorphus, 10918.
tympanus Lonnberg and Andersson, Sphenomorphus, 1913.
undulatus Peters and Doria, Sphenomorphus, 1878.
umcolor Harlan, Scincella, 1825.

unilineatus de Rooij, Sphenomorphus, 1915.
vandenburghi Schmidt, Scincella, 1927.
variabilis Bavay, Lampropholis, 1869.
variegatus Peters, Sphenomorphus, 1867.
verreauxii Duméril, Saiphos, 1851.
vertebrale de Vis, Carlia, 1888.

victoriana Shreve, Scincella, 1940.
vigintiserius Sjostedt, Sphenomorphus, 1897.

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL) La7)

virens Peters, Lipinia, 1881.

virgatus Garman, Cryptoblepharus, 1901.
viridipunctata Lesson, Dasta, 1830.

vittata Ederling, Dasia, 1864.

vittigera Boulenger, Scincella, 18094.

vivax de Vis, Carlia, 1884.

voeltzkoui Sternfeld, Cryptoblepharus, 1918.
vosmaerit Gray, Riopa, 1839.

vulcamia Girard, Scincella, 1857.

vyneri Shelford, Dasia, 1905.

wahlbergti Smith, Ablepharus, 1849.

walkert Boulenger, Nodorha, 1801.

webert Schmidt, Leptosiaphos, 1943.
weeksae Kinghorn, Leiolopisma, 1928.
wernerit Vogt, Emoia, 1912.

wetariensis Mertens, Sphenomorphus, 1928.
whitneyt Burt, Emoia, 1930.

wilkinst Parker, Nodorha, 1926.

wolfi Sternfeld, Otosaurus, 1918.

wolfi Sternfeld, Sphenomorphus, 1918.
wollastoni Boulenger, Sphenomorphus, 1914.
woodfordi Boulenger, Sphenomorphus, 1887.
woodwardi Lucas and Frost, Hemiergis, 1894.
yunnanensis Boulenger, Tropidophorus, 1887.

BIBLIOGRAPHY

The bibliography of the Lygosominae is enormous, scattered as it
is over hundreds of miscellaneous papers, monographs, exploration re-
ports, and general zoological treatises which have appeared during
the last 150 years. In consequence, rather than being a compendium
of this literature the present bibliography includes only certain stan-
dard works and geographic studies to which the interested student is
referred for more specific literature treating forms of particular
interest.

Bargour, THOMAS.
1912. A contribution to the zoogeography of the East Indian islands. Mem.
Mus. Comp. Zool., vol. 44, pp. 1-203, pls. 1-8.
1921. Reptiles and amphibians from the British Solomon Islands. Proc.
New England Zool. Club, vol. 7, pp. 91-122, pls. 2-4.
BouLENGER, GEORGE A.
1887. Catalogue of the lizards in the British Museum. 2d ed., xii + 575 pp.,
40 pls. London.
Burt, CHARLES E., and Burt, May D.
1932. Herpetological results of the Whitney South Sea Expedition.
VI. Bull. Amer. Mus. Nat. Hist., vol. 63, pp. 461-597, figs. 1-38.
Cope, Epwarp D.
1864. Contributions to the herpetology of tropical America. Proc. Acad.
Nat. Sci. Philadelphia, vol. 3, pp. 166-176; vol. 4, pp. 177-181.

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 33

1892. On degenerate types of scapular and pelvic arches in the Lacertilia.
Journ. Morph., vol. 7, pp. 223-244, pl. 13.

1900. Crocodilians, lizards, and snakes of North America. Ann. Rep. U. S.
Nat. Mus. for 1898, pp. 153-1204, figs. 1-347, pls. 1-36.

DumEriL, A. M. C., and Brsron, G.
1839. Erpétologie générale. . . . 5 vols., viii +854 pp. Paris.
DumeErtL, A., and Bocourt, Marte-Frrmin.

1881. Etudes sur les reptiles et les batraciens. Mission Scientifique au
Mexique et dans l’Amérique Centrale, vol. 3, No. 7, pp. 441-488,
pls. 22e-22)j.

DumeriL, A. M. C., and Dumérit, M. A.
1851. Catalogue méthodique de la collection des reptiles du Muséum d’His-
toire Naturelle de Paris. iv +224 pp. Paris.
FitrziNncer, LEopotp J.
1826. Neue Classification der Reptilien. 66 pp., 1 pl. Wien.
1843. Systema reptilium. . .. 106 pp. Vindobonae.
FITZSIMONS, VIVIAN F.

1943. The lizards of South Africa. Transvaal Mus. Mem., vol. 1,

xv + 528 pp., 384 figs., 24 pls., map.
Gray, JoHN E.

1831. A synopsis of the species of the class Reptilia. (App., pp. I-100) in
Cuvier, The Animal Kingdom. Class Reptilia. Griffith ed., 480 +
100 pp., 54 pls. London.

1845. Catalogue of the specimens of lizards in the collection of the British
Museum. xxvii + 289 pp. London.

GuntuHeEr, Apert C. L. G.
1864. The reptiles of British India. xxvii + 452 pp., pls. 1-26. London.
HorrMann, C. K.

1881. Klassen und Ordungen des Thier-Reichs, etc. Bd. 6, Abt. 3. Rep-

tilien, Lief. 18-21, pp. 443-1399, pls. 49-107. Leipzig and Heidelberg.
LovERIDGE, ARTHUR.

1928. Field notes on vertebrates collected by the Smithsonian-Chrysler
East African expedition. Proc. U. S. Nat. Mus., vol. 73, No. 17,
pp. 1-60, pls. 1-4.

1933. Reports on the scientific results of an expedition to the southwestern
highlands of Tanganyika Territory. VII. Herpetology. Bull.
Mus. Comp. Zool., vol. 74, No. 7, pp. 195-416, 3 pls.

1933. New scincid lizards of the genera Sphenomorphus, Rhodona and
Lygosoma from Australia. Occ. Pap. Boston Soc. Nat. Hist.,
vol. 8, pp. 95-100.

1934. Australian reptiles in the Museum of Comparative Zoology, Cam-
bridge, Massachusetts. Bull. Mus. Comp. Zool., vol. 77, No. 6,
Pp. 243-383, 1 pl.

1942. Scientific results of a fourth expedition to forested areas in East
and Central Africa. IV. Reptiles. Bull. Mus. Comp. Zool., vol. 91,
No. 4, pp. 237-373, pls. 1-6.

1948. New Guinean reptiles and amphibians in the Museum of Comparative
Zoology and United States National Museum. Bull. Mus. Comp
Zool., vol. 101, No. 2, pp. 305-430.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

MERTENS, ROBERT.
1928. Neue Inselrassen von Cryptoblepharus boutoniit (Desjardin). Zool.
Anz., vol. 78, pp. 82-89.
1929. Die Rassen des Smaragdskinkes Dasia smaragdinum Lesson. Zool.
Anz., vol. 84, pp. 209-220.
1930. Die Amphibien und Reptilien der Inseln Bali, Lombok, Sumbawa und
Flores. Abh. Senck. nat. Ges., vol. 42, pp. 117-344, pls. 1-4.
1931. Ablepharus boutonit (Desjardin) und seine geographische Variation.
Zool. Jahrb., vol. 61, pp. 63-210, pls. 2-4.
MiTTLEMAN, M. B.
1950. The generic status of Scincus lateralis Say, 1823. Herpetologica,
vol. 6, No. 2, pp. 17-20.
Peters, W., and Dorta, G.
1878. Catalogo dei rettili e dei batraci raccolti da O. Beccari, L. M.
D’Albertis e A. A. Bruijn nella sotto-regione Austro-Malese. Ann.
Mus. Civ. Stor. Nat. Geneva, vol. 13, pp. 323-450, pls. I-7.
Roory, NELLY DE.
1915. The reptiles of the Indo-Australian Archipelago. I. Lacertilia,
Chelonia, Emydosauria. xiv + 384 pp., 132 figs. Leiden.
Scumipt, Kart P.
1927. The reptiles of Hainan. Bull. Amer. Mus. Nat. Hist., vol. 54,
Pp. 395-465, 17 figs., pl. 27.
1927. Notes on Chinese reptiles. Bull. Amer. Mus. Nat. Hist., vol. 54,
pp. 467-551, 22 figs., pls. 28-30.
1943. Amphibians and reptiles from the Sudan. Zool. Ser. Field Mus. Nat.
Hist., vol. 24, No. 20, pp. 331-338, 1 fig.
SmitH, Hopart M., and TayLor, Epwarp H.
1950. An annotated checklist and key to the reptiles of Mexico, exclusive
of the snakes. U. S. Nat. Mus. Bull. 199, v + 253 pp.
SmitH, Matcotm A.
1927. Contributions to the herpetology of the Indo-Australian region. Proc.
Zool. Soc. London, 1927, pp. 199-225, 2 pls.
1935. The fauna of British India... . Reptilia and Amphibia. 2, Sauria.
xiii + 440 pp., I pl., 93 figs. London.
1937. A review of the genus Lygosoma (Scincidae: Reptilia) and its allies.
Rec. Indian Mus., vol. 39, No. 3, pp. 213-234, figs. 1-3.
STEJNEGER, LEONHARD.
1907. The herpetology of Japan and adjacent territory. U. S. Nat. Mus.
Bull. 58, xx + 577 pp., 400 figs., 35 pls.
1925. Chinese amphibians and reptiles in the United States National Mu-
seum. Proc. U. S. Nat. Mus., vol. 66, No. 25, pp. 1-115, 6 figs.
STERNFELD, RICHARD.
1918. Neue Schlangen und Ejidechsen aus Zentralaustralien. Mitt. Senck.
Ges., vol. 1, pp. 76-82.
1925. Beitrage zur Herpetologie Inner-Australiens. Abh. Senck. nat.
Ges., vol. 38, pp. 221-251.
Stuart, LAURENCE C.
1940. Notes on the “Lampropholis” group of Middle American Lygosoma
(Scincidae) with descriptions of two new forms. Occ. Pap. Mus.
Zool., Univ. Michigan, vol. 421, pp. 1-16, I fig.

NO. 17 LIZARDS OF SUBFAMILY LYGOSOMINAE—MITTLEMAN 35

TAyYLor, Epwarp H.
1922. The lizards of the Philippine Islands. Bur. Sci. Manila Publ. 17,
269 pp., 53 figs., 23 pls.
Van DENBURGH, JOHN.
1912. Concerning certain species of reptiles and amphibians from China,
Japan, the Loo Choo Islands and Formosa. Proc. California
Acad. Sci., vol. 4, No. 3, pp. 187-258.
Waite, Epcar R.
1929. The reptiles and amphibians of South Australia. 270 pp., 192 figs.
Adelaide.
Zietz, F. R.
1920. Catalogue of Australian lizards. Rec. South Austral. Mus., vol. 1,
pp. 181-228.

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Charles QD. and Mary Waux Halcott
Research Fund

|| | THE LOWER EOCENE KNIGHT
|| FORMATION OF WESTERN WYOMING
‘|| AND ITS MAMMALIAN FAUNAS

(WirH 11 Pirates)

BY
C. LEWIS GAZIN

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Curator, Divison of- Vertebrate Paleontology
United States National Museum

maven TO?

(Pusiication 4097)

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(eae "PUBLISHED BY THE SMITHSONIAN INSTITUTION
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FORMATION OF WESTERN WYOMING
AND ITS MAMMALIAN FAUNAS

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 117, NUMBER 18
(Enp oF VoLuME)

Charles D. and Mary Waux Walcott
Research Fund

fig LOWER EOCENE KNIGHT
FORMATION OF WESTERN WYOMING
AND ITS MAMMALIAN FAUNAS

(Wir 11 Pirates)

BY
C. LEWIS GAZIN

Curator, Divison of Vertebrate Paleontology
United States National Museum

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(Pus.ication 4097)

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PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 9, 1952

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BALTIMORE, MD., U. 8. A.

CONTENTS

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

ILLUSTRATIONS

PLATES

(All plates following p. 82.)

. Marsupials, insectivore, and primates from the Knight lower Eocene.
. Insectivore and taeniodont from the Knight lower Eocene.

. Taeniodont foot from the Knight lower Eocene.

. Creodonts from the Knight lower Eocene.

. Meniscotherium from the Knight lower Eocene.

. Coryphodon from the Knight lower Eocene.

. Lambdotheriwm from the Knight lower Eocene.

. Artiodactyles from the Knight lower Eocene.
FIGURES

. Map of southwestern Wyoming and portions of adjacent States....... 5
. Pentapassalus pearcet, new genus and SpeCieS..........eeseccceccceace 33
. Pentapassalus pearcei, new genus and specieS.............eeeecceseees 40
. Pentapassalus pearcet, new genus and species..........-...-e2- eee {Race sAL
« Lillomys ‘sentor;, WHEW ,SPCCIES., «jsa's's «isis a: «a0 stew sly se +d RAE eR ee ae.)
. Vaverravus: Intosus, New ‘SPeCleS. «0.04 sd0s's.00tosbasw Ckew ks ORR 56

Charles BD. and Mary Waux Walcott Reseach Fund

THE LOWER EOCENE KNIGHT FORMATION OF
WESTERN WYOMING AND ITS
MAMMALIAN FAUNAS

By C. LEWIS GAZIN

Curator, Division of Vertebrate Paleontology
United States National Musewmn

(WitTH 11 PLATEs)

INTRODUCTION

The variegated beds of the Wasatch group in southwestern Wyo-
ming have been of much interest to both geologists and paleontologists
since their description by Hayden in 1869. To vertebrate paleontolo-
gists significance lies largely in their having yielded the first known
lower Eocene mammals from North America. Unfortunately, how-
ever, the term “Wasatch” was too broadly defined for precise geologic
usage and included strata of more than one geologic age. In conse-
quence of this, Veatch in 1907 redefined the term as a group and
divided it in ascending order into the Almy, Fowkes, and Knight
formations. The last-named formation, including the fossiliferous
lower Eocene horizons, is the primary concern of this report.

Investigation by the Smithsonian Institution of the mammalian
faunas of the Knight formation was largely an outgrowth of the
discovery by U. S. Geological Survey parties of Paleocene vertebrate
remains in underlying Almy deposits (Gazin, 1942). Collections by
J. B. Reeside, Jr., W. W. Rubey, and B. N. Moore in 1936 and by
Rubey and John Rogers in 1939 led us in 1941 to a careful examina-
tion of the Almy beds as exposed on La Barge Creek in Lincoln
County, Wyo., and to a somewhat desultory search of the nearby
Knight exposures along Green River in Sublette County. Examina-
tion of the two sets of beds was renewed in 1948, and an intensive
search of the Knight exposures was made along the east side of Green
River from the vicinity of the “Three Bridges” southeast of Big
Piney to 10 or 11 miles south on Fogarty Draw. In 1949 search was
extended to both sides of the Green River, on the west side as far
south as La Barge, and to about 12 miles north of Big Piney. In 1951

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 117, NO. 18

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

re-examination was made of all the more profitable localities and in-
vestigation was carried upward into the New Fork member, a tongue
of Wasatchian strata interfingering with the lowermost Green River
beds in the same area.

Collections made from the upper Green River basin, together with
materials from the Knight beds in the vicinity of Fossil and those
of the earlier collections from the Knight on Bear River, result in our
now recognizing at least four horizons for fossil mammals in pre-
Bridgerian Tertiary of southwestern Wyoming. These include the
Clarkforkian Paleocene stage represented by mammalian remains
from the Almy on La Barge Creek, equivalent to part of the original
Wasatch; a Lysite or about middle-Wasatchian stage represented by
collections from the type locality for the Knight, and probably the
Knight at Fossil; and apparently two horizons of general Lost Cabin
equivalence supported by the collections from the upper Knight beds
and New Fork beds along the Green River in Sublette County. The
lower Wasatchian or Gray Bull has not been certainly recognized,
except in the adjacent Hoback basin (see Dorr, 1952).

ACKNOWLEDGMENTS

In addition to the above-named members of the U. S. Geological
Survey, I wish to acknowledge helpful information given me by
Dr. G. E. Lewis, who made a small collection from Knight beds near
the Green River in 1947 while working with Dr. Rubey on the Big
Piney Quadrangle. George Shea, of Billings, Mont., very graciously
aided in locating for me materials which he and G. E. Lewis ob-
served in 1947. Acknowledgment is also made of the courtesies ex-
tended in connection with my work in 1951 by the geologists of the
General Petroleum Co., the Mountain Fuel and Supply Co., and the
Shell Oil Co. doing exploratory work in the same area. My field
assistants in 1941 included G. F. Sternberg and F. L. Pearce. In
1948 I was aided by my wife, Elisabeth, and son, Chester Gazin ;
both accompanied F. L. Pearce and myself in 1949.

In the faunal study I am indebted to Dr. Glenn L. Jepsen for per-
mitting me to examine and describe Knight materials in the Princeton
University collections and make comparisons with type and other speci-
mens there. Drs. G. G. Simpson, E. H. Colbert, and Bobb Schaeffer
extended to me facilities of the American Museum of Natural History
and permitted me to study Knight materials from the type locality and
various type specimens of lower Eocene mammals in the American
Museum collections.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 3

The drawings depicting new forms in figures 2-6 were prepared by
William D. Crockett. The photographs of specimens in plates 1, 2
(fig. 1), 5-8, 10, and 11 were made through the kindness of Dr.
G. Arthur Cooper. Photographs in text figure 1, plate 2 (figs. 2-5),
and plates 3, 4, and 9 were made by the photographic laboratory of the
U. S. National Museum.

HISTORY OF INVESTIGATION

Special historic interest, as has been noted, is involved in considera-
tion of the Knight formation and its faunas. Fossil mammalian re-
mains from exposures of this formation along Bear River, near
Evanston, Wyo., are apparently the first to be described from the
lower Eocene of North America. From here Hayden, or Cleburne
according to Marsh (1893, p. 321), in 1871 secured the remains that
Cope (1872a) described as “Bathmodon” radians and “Bathmodon”
semicinctus, and part of the material later described as “Bathmodon”
latipes. In 1872 Cope, assisted by Garman, and while working for
the Hayden Survey, collected the type of Hyracotherium vasacciense
as well as materials that became the types of various turtles (1872b).
It is of interest that in 1872 Cope’s travels took him to the mouth of
La Barge Creek in the general area of upper Knight beds that produced
the materials forming the basis of this investigation. It is unlikely,
however, that he made any collection there. The same year he ob-
tained from the vicinity of Black Buttes, east of the Rock Springs up-
lift, teeth from two badly decomposed Coryphodon skulls that he
named (1873b) “Metalophodon” armatus. Apparently again in 1873
Cope went over the exposures along Bear River, southeast of Evans-
ton, and collected the type materials of Hyracotherium index and
Phenacodus primaevus (1873a), and additional material of Corypho-
don radians and Coryphodon latipes.

Cope’s investigation in the Knight ended with his transfer of in-
terest to the lower Eocene of New Mexico and our next record of
activity in these beds is Marsh’s description of Coryphodon hamatus
in 1877. This was collected in about 1874, from exposures on Bear
River, about “35 miles west of Bridger,” by R. Veltman of Evanston,
Wyo., formerly a storekeeper at Fort Bridger. Veltman first wrote *
Marsh about having discovered large bones on Bear River in June of
1871. Marsh apparently did not follow up this information until
Veltman wrote again in November of 1874, at which time he reported

1Information kindly furnished me by Dr. J. T. Gregory from the file of
Marsh’s correspondence.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. TL7

finding a skull and other bones of a very large animal, offering them
for sale. These were evidently of the Coryphodon hamatus skeleton
that Marsh eventually obtained. About the middle of the same decade
Marsh acquired material of Hyracotherium, from the vicinity of
Black Buttes, that in 1876 he named Eohippus pernix. William Cle-
burne, of the Union Pacific Railroad, in 1875 collected the skull and
jaws that Thorpe (1934) described as Meniscotherium robustum.
The latter was found in a cut on the old grade of the railroad about
2 miles west of Aspen (old location).

I have no further information on early paleontological exploration
in these beds until 1906, when Granger and Miller collected about a
dozen specimens of small mammals approximately 200 feet up on the
bluffs of Bear River south of Knight station and in the vicinity of
the railroad bridge. These are specifically mentioned by Granger in
1914 (p. 203) and regarded as Lysite in age.

More recent active interest in the Knight was recorded by Bonillas
when, in 1936, with his description of the dentition of Lambdotherium
from deposits near La Barge, Wyo., he mentioned the field exploration
of the California Institute of Technology in these beds and cited
briefly the fauna obtained. Their collecting was done largely in the
vicinity of La Barge and included search on both sides of the Green
River. Field work by parties from Princeton University in the Knight
formation resulted in the discovery in 1939 and 1940 of fragmentary
mammalian remains in Knight beds 40 to 100 feet below Green River
strata about 3 miles northwest of Fossil, Wyo. Also, a jaw of Am-
bloctonus as well as an isolated tooth of Oxyaena were found by
them southeast of La Barge in 1941. These, together with the above-
noted finds of the U. S. Geological Survey and the more extensive
exploration and collecting by the Smithsonian Institution, complete
the history of paleontological work in the Knight of the Fossil and
upper Green River basins, as far as it is known to me.

OCCURRENCE AND PRESERVATION OF MATERIAL

For practical purposes the Knight collections may be regarded as
coming from three general localities or areas of exposure in the
westernmost part of the State: one in the northwestern part of the
upper Green River or Bridger basin and two in the adjacent, small
but elongate structural basin to the west which includes the type
section. In addition to these are a few isolated occurrences either at
remote localities tentatively regarded as Knight or in exposures sep-
arated from the general localities by significant faulting.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 5

Wie:
»))
ig? Bs

Fic. 1.—Map of southwestern Wyoming and portions of adjacent States show-
ing Eocene sedimentary basins, with fossil localities in Knight formation num-
bered as follows: 1, Knight station, type Knight; 2, Fossil; 3, La Barge—Big
Piney; 4, New Fork; 5, Rock Springs; 6, Vermilion Creek; 7, Black Buttes ;
8, Dad-Baggs; 9, Great Divide basin; 10, Red Desert. Map reproduced from
Osborn, U. S. Geol. Surv. Monogr. 55, figs. 9, 49, 1929.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

Historically, the more important of the localities would be the ex-
posures along Bear River, in general between Evanston and Knight
station. Granger’s collection came mostly from the bluffs south of
Knight station, and the only detailed statement made by Cope (1872b,
p. 473) regarding his collection refers to “upper red and white strata
on the bluffs eleven miles S. E. of Evanston, or near the bend of Bear
River.” Except for the Coryphodon hamatus skeleton collected for
Marsh, the materials so far recovered from this area have been very
fragmentary. Those collected by Granger and Miller occurred in a
yellowish or buff sandstone, somewhat conglomeratic, hence conditions
of deposition were not the most favorable for preservation of the
smaller mammals. The bone portion of the jaws of the smaller
mammals is very poorly preserved and only the teeth are in reasonably
good condition.

The exposures near Fossil, Wyo., from which Princeton obtained
a collection, are about 45 miles due north of Knight station but in the
same basin of deposition. The Princeton collection was made in
variegated beds from 40 to 100 feet beneath the Green River forma-
tion about 3 miles north of Fossil. However, a jaw of Phenacodus
was obtained by Dr. Dunkle for the Smithsonian Institution in gray
exposures beneath the red beds just 3 miles to the east of Fossil.
The materials from near Fossil are also fragmentary but the bone
has more the appearance of that which occurs in the vicinity of La
Barge and Big Piney.

The most prolific collecting ground in beds considered to be a part
of the Knight formation are to be found in the vicinity of the Green
River in Sublette County—from La Barge, approximately 40 miles
northeast of Fossil, to a point about 12 miles north of Big Piney, Wyo.
Fossil materials of this area are found in many places on both sides
of the Green River; however, certain small patches or coves of ex-
posures have proven much more productive than others. Materials
here were encountered at various levels from as near as about 40 feet
below Green River strata, to possibly 100 feet lower. A collection
was also obtained from the New Fork tongue of Wasatchian beds
in the lower part of the Green River section along Alkali Creek and
New Fork, tributaries of the Green River to the east and southeast
of Big Piney.

The Smithsonian collections, which are almost exclusively from the
La Barge—Big Piney area, number a little under 600 and include ma-
terials in various states of preservation and completeness. Excellent

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 7

skulls were found of some forms, showing varying amounts of dis-
tortion, but less in general than exhibited by materials from lower
Eocene deposits in the Big Horn, Wind River, and San Juan basins.
Not all specimens are free of an iron-oxide coating, but the prevalence
of this condition is far less than in the Big Horn basin Willwood ma-
terials. No one horizon or lithology can be regarded as exclusively
fossiliferous, except locally and for short distances, although certain
dark reddish to purplish zones to the east of the Green River have
been relatively productive, and a soft blue-gray shale at certain places
on the west side between La Barge and Big Piney. A rather profit-
able concentration of materials was encountered about 12 miles north
of Big Piney in reddish-buff layers and gray sandy shale.

Isolated occurrences include the Aspen locality for the type of
Mentscotherium robustum (in the Green River basin of deposition
and hence not to be confused with the type Knight), and the occur-
rence of Coryphodon and Hyracotherium near the upper reaches of
Dry Piney Creek, referred to in 1942. In the eastern part of the
Bridger basin, around the Rock Springs uplift, isolated occurrences
presumably in the same formation include a Meniscotherium robustum
skull collected by Roland W. Brown just southwest of Rock Springs ;
a small collection of fragmentary remains that I made from near the
southern extremity of the uplift on a tributary of Vermilion Creek ;
and the Black Buttes occurrence to the east of the uplift. Still more
remote, but nevertheless in the same depositional basin, may be men-
tioned the occurrences on the eastern margin of the Washakie basin
beneath the Tipton tongue of Green River between Dad and Baggs,
referred to in Wood el al. (1941, p. 18), and from northeast of Cres-
ton made by a U. S. Geological Survey party in 1907.

THE KNIGHT FAUNAS

The following tabulation pertains to the principal occurrences of
fossil mammals in the Knight formation as it is exposed in the upper
Green River basin (the La Barge and New Fork faunas) and in the
adjacent small structural basin west of the Oyster Ridge which in-
cludes the type Knight and the occurrence at Fossil, Wyo. The figures
used refer to the number of specimens encountered, giving an indica-
tion of extent of materials upon which identifications are based, and
a suggestion of the relative abundance of various forms in the faunas.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Knight La Barge- New
MARSUPIALIA : station Fossil Big Piney Fork

Peratherium edwardi, new species....... 2

Peratherium chesteri, new species....... I
INSECTIVORA :

Diacodon pineyensis, new species........ 3

Diacodon, cf. alticuspis Cope........... : I

Cynodontomys knightensis, new species. . 13

Cinodontomnys. Species she ea ssn oaeiices I
TILLODONTIA :

Esthonyx, ci. acutidens Cope............ 3

EEStN ONY NSDECICS Raa eee eee I I
PRIMATES :

ely COdUS a ASNECleSh Eee Eee se eae I

Notharctus limosus, new species......... 24

Notharctus venticolus Osborn........... 6

Absarokius noctivagus Matthew.........

Paratetonius? sublettensis, new species... I
TAENIODONTA :

undetermined stylinodont .............. ; 4
EDENTATA:

Pentapassalus pearcei, new genus and

SPECICSiy Patire i Ha eth eee yep veetey tee

RODENTIA :

Paramys, cf. buccatus * (Cope).......... I

Paramys, ci. copei* Loomis.............

Sciuravus, possibly S. depressus Loomis. .

Tillomys semor, new species...........;
CARNIVORA :

Thryptacodon, near T. antiquus Matthew. 2

Pachyaena? Species ps aceon a see ee ee 2

ORNGENG SPECIES A a. davis aN leremtalaeh &

Ambloctonus, cf. major Denison.........

Prolimnocyon clisabethae, new species...

Sinopa vulpecula Matthew.......... aint

Sinopa, cf. strenua (Cope).........0.005

Didymictis altidens Cope..............- ?

Viverravus lutosus, new species.........

Uintacyon asodes, new species.........-.

Miuacis, cf. latidens Matthew....... Saas

Vulpavus asius, new SpecieS..........-- é
CoNDYLARTHRA :

Hyopsodus wortmani Osborn.......... ; 29 I

Hyopsodus mentalis (Cope)...........- 76

Hyopsodus brownt Loomis........... Stuy!

Phenacodus primaevus Cope.........0+ I 1 *

Meniscotherium robustum Thorpe....... 93

Meniscotherium, cf. chamense Cope...... 3

See ON

La)
AW FH OPH MNW HH

* Paramys, cf. buccatus in Knight is Paramys excavatus Loomis; and Paramys, cf. copei
Loomis includes P. copei, P. major, and a new species, as determined by A. E. Wood.
** Recorded from a horizon stratigraphically lower than other forms cited in this fauna.

no. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 9

PANTODONTA: pied Fossil Bae rae
Coryphodon radians (Cope)............ Xf Chats
Coryphodon semicinctus (Cope)........ I
Coryphodon latipes (Cope)............. I
Coryphodon hamatus Marsh............ I
GOrupnodon: SPECIES... oo sc cose oe aes « I

DINOCERATA :

Ci. Bathyopsis fissidens Cope............ 2

PERISSODACTYLA :

Hyracotherium index (Cope)........... 2 I 35
Hyracotherium vasacciense (Cope)..... 4 67
Hyracotherium, cf. venticolum Cope..... I 4 3
Hyracotherium, possibly H. cristatum

WiOnINate oases eee Seat oceat ome e io. I
Lambdotherium popoagicum Cope....... 93 II
Heptodon, cf. ventorum (Cope).........- I 30
lie PLOMGIAaSPCCIES: Mirsersieiaiess inis:s)soisie aes os I
TA KOGHSUSH aS PECCICS a cers alate s(cvagels.e- 015/510, sieleis I

ARTIODACTYLA :

Bunophorus, ci. macropternus (Cope)... 7)
Diacodexis, near D. secans (Cope)...... I
IDOE T IS, GUAGE Gooodd sobundondnes oc De

He-xacodus pelodes, new genus and species. II
Hexacodus uintensis, new species........ I

{+ Number of specimens not known and should include material of at least a part of those
species listed below which are probably synonyms.
Specimens not seen.

CORRELATION AND AGE OF FAUNAS

Within the formation that may properly be called Knight are
clearly three horizons that can be distinguished on the basis of fossil
mammalian remains, including the New Fork tongue of Wasatchian
material interfingering in the lower part of the Green River series.
The lower of these is nearest to Lysite in age and the upper are two
distinct zones of Lost Cabin age separated locally by the Fontenelle
tongue of Green River. In the absence of evidence of any lower
Wasatchian or Gray Bull horizon, except in the Hoback basin (see
Dorr, 1952), the Knight may well be regarded as about equivalent
to the Wind River series.

Knight station —Critical examination of the materials derived from
the type Knight beds along Bear River lead me to support Granger’s
(1914, p. 203) conclusion that the horizon there represented is about
equivalent to Lysite. The Cynodontomys material, as incompletely
known, does not appear to be more indicative of a Lysite species

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

than of the later form now known to characterize the La Barge fauna;
however, Hyopsodus browni is highly suggestive of Lysite and the
Heptodon material would not appear to be older. The relative fre-
quency with which Coryphodon remains have been encountered would
appear to have no significance, and although not recorded in the type
Lysite they are found in Lost Cabin beds of both Wind River and
Green River basins, as well as in the Gray Bull, so that their middle
Wasatchian or Lysite representation may well be here. Among the
14 to 16 forms recognized in the fauna, characteristic or peculiarly
Gray Bull types are missing, particularly such a dominant form as
Homogalax. In a like manner, the absence of any representation of
Lost Cabin forms as Meniscotherium and Lambdotherium is equally
suggestive. In support of this it should be noted that, with regard
to the Lost Cabin equivalent of the Knight in the upper Green River
basin, almost a third of the specimens encountered are of one or the
other latter forms. In a cursory examination of the upper Knight
the percentage of these forms would, because of their size, appear
to be even much higher.

Fossil_—The rather limited fauna represented by the small collection
from near Fossil, Wyo., is less clearly diagnostic with respect to the
horizon represented than is the Bear River Knight. Comparison with
horizons of the Wind River and Big Horn basins leaves much to be
desired; nevertheless, its closest affinity is with the type Knight.
Heptodon, taken into consideration with other similarities to the
fauna from the Knight on Bear River, tend to restrict comparison to
only the middle and perhaps upper Wasatchian levels. On the other:
hand the species of Diacodon and Hexacodus represented are dis-
tinctly not those found in the Lost Cabin equivalent of the Knight in
the Green River basin, and Hexacodus is not among the rarer forms
represented in the latter beds. Phenacodus primaevus, though found
at a somewhat lower level than other materials from the Fossil area,
is significantly absent from the La Barge fauna. It must be noted,
however, the situation with respect to Phenacodus is reversed in the
Wind River basin. Individually, the elements of the Fossil fauna
are inconclusive, but collectively there is a strong presumption toward
the Lysite level as that level is indicated by the Knight fauna obtained
from the Bear River section.

La Barge.—There would appear to be no doubt of the general
equivalance of the La Barge fauna to the Lost Cabin of the Wind

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN ial

River basin. There is a specific difference between them in many
of the genera mutually represented, but this may be largely environ-
mental, or possibly of some small time significance if within a rapidly
evolving group. Such a time difference, however, would not be of the
magnitude of that between Lysite and Lost Cabin. The Lost Cabin
equivalance is indicated by the presence of such genera as Notharctus,
Meniscotherium, and Lambdotherium which first appear, or reach
the indicated development, in the Lost Cabin horizon of the Wyoming
lower Eocene; by suclr species as Esthony-x, cf. acutidens; Notharctus
venticolus; Absarokius noctivagus; Ambloctonus, cf. major; Sinopa
vulpecula; Muacis, cf. latidens; Hyopsodus mentalis; Hyopsodus
wortmam,; Hyracotherium, cf. venticolum; and Lambdotherium popo-
agicum which are close or identical to such species more or less
characteristic of Lost Cabin in the Wind River or Big Horn basins;
and by the developmental stage reached in such species as Cynodon-
tomys knightensis, Notharctus limosus, and Prolimnocyon elisabethae
which are otherwise distinct from related Lost Cabin species.

Suggestive of perhaps a slightly earlier stage than type Lost Cabin,
although as noted above these differences may be no more than en-
vironmental in nature, are the following: The smaller size, though
equivalent dental development, of Cynodontomys knightensis in com-
parison with Cynodontomys scottianus in a sequence of species in-
creasing in size; the smaller size and relatively smaller premolars in
Notharctus limosus than in N. nunienus; the weaker hypocone in
upper molars of Notharctus venticolus of the La Barge fauna; the
smaller size (of questionable significance) of Diacodon pineyensis ;
Esthonyx, cf. acutidens; Absarokius noctivagus; Prolimnocyon elisa-
bethae; Sinopa vulpecula; Didymictis altidens; Viverravus lutosus;
and Vulpavus asius in comparison with the same or equivalent Lost
Cabin species ; and the presence of Paratetonius?, Thryptacodon, and
Hyracotherium index. The above would appear to be of perhaps
trifling importance in consideration of relative ages, but on the other
hand there is even less to suggest a later stage than type Lost Cabin,
although such might be interpreted from the presence of Sciuravus,
Tillomys, and Hexacodus. The evidence for a slightly older age
afforded by Meniscotherium is perhaps the most interesting, inasmuch
as the large MW. robustum so far not found outside of the Green River
basin occurs stratigraphically below Meniscotherium, cf. chamense
of the New Fork tongue. Only the smaller species, regarded as
M. chamense, is recorded from the Lost Cabin beds of the Wind
River basin.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

The following tabulation is a comparison of genera now known in
the La Barge and Lost Cabin faunas:

Lost Lost
La Barge Cabin La Barge Cabin

Peratherium ....... x IPyotopsaiis seme ces xX
IDWeCO CIO! dog Aan 60.4 xX x Ambloctonus ....... xt x
Parietops \. ecto Xx Prolimnocyon ...... x x
Didelphodus ........ Xx Sanofi) Wath hae ok x x
Palagosinopa ......: Xx LGV UCTS waite esas 3 x x
Cynodontomys ..... x x VAGCLF GUUS, seis, s4 x x
ESiHOnV es Vener eon eee D4 D4 Omiacyon eee ces Xx

Notharctus 050.0. . xX x MEGICLEN Cee ad aeat x xX
EOUCeina onsen ek we Xx VaUpQUUs 22a Scat x xX
Shoshonius......... Xx Phenacodus ........ xX
Ti CLOWUES ce voces sine xX FSCEOCION: Seale: a.0 «rie! seh x
Paratetonius? ...... x Mentscotherium .... X x
Absarakwms (21453. Xx Xx Hyopsodus -}...-.:: x x
Stylniodon s. Sea. cf. xX Coryphodon ........ xX xX
palaeanodont ....... x Bathyopsis .......+5 xX
TUbUlOdOn, 5 Fs occ 0.0 » xX Hyracotherium ..... X x
Pentapassalus ...... xX Lambdotherium . xX xX
POFAMY Socrates xX x Eoniagnops 3... <<< ».
SCH AUUS I Te, Poe xX Hepiodow ss c-6.: xX xX
disllomy sapere. x Fy rachyus con voce: xX
MSS OP Sind svete tay: xX Bunophorus ........ xX x
Thryptacodon ...... Xx Digcodexis 5. s.cnteee x xX
OV UeHOn arereere X x IIL BOUTS n56d0000 xX

New Fork.—The New Fork tongue, separated by Green River
beds called the Fontenelle tongue by Donovan (1950) from the
Knight along the Green River, has yielded a faunal representation
which compares equally well with that of the Lost Cabin, but includes
forms not found in the La Barge fauna. The Lost Cabin age is here
indicated by Ambloctonus, cf. major; Hyopsodus wortmani; Menis-
cotherium, cf. chamense; cf. Bathyopsis fissidens; Hyracotherium, cf.
venticolum; and Lambdotherium popoagicum. The significant dif-
ferences to be noted from the La Barge fauna are in (1) the species
of Meniscotherium represented, (2) the presence of Bathyopsis? in-
stead of Coryphodon, and (3) the appearance of Hyrachyus. I suspect
that, were the New Fork better known, Eotitanops would be repre-
sented, also Trogosus, which is known from the Cathedral Bluff
tongue in the northwestern part of the Red Desert.

The most significant feature of the discovery of the New Fork
fauna is the clear demonstration that the lower beds of the Green
River are actually Lost Cabin Wasatchian in age and that the Lost

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN Ae)

Cabin fauna can be largely broken up into two stages. Tentatively
the lower stage may be regarded as characterized by Meniscotherium
robustum and Coryphodon and the upper stage by Meniscotherium
chamense, Hyrachyus, species, and possibly Bathyopsis. Osborn early
regarded the Lost Cabin as comprising two stages, but believed them
to be distinguished by Lambdotherium and Eotitanops, respectively ;
however, Lambdotherium is here found in both levels. Much more of
the New Fork fauna needs to be known for a more complete under-
standing of Lost Cabin zones. This may come when the Cathedral
Bluff fauna currently under study by William Morris is placed on
record.

ISOLATED OCCURRENCES TO THE EAST OF THE
BRIDGER BASIN

Not included in the foregoing tabulation and correlations, and for
the most part omitted from consideration in the systematic treatment
of the faunas, are the scattered occurrences in the eastern part of the
Bridger basin, around the Rock Springs uplift, in the eastern part
of the Washakie and Great Divide basins, and in the Red Desert.
It is not certainly known but entirely probable that the formation rep-
resented by these occurrences is also Knight. An examination of the
geologic map of Wyoming leads to the conclusion that the Eocene
basin of deposition is the same for these deposits as for those in the
vicinity of La Barge and Big Piney, and that the formation is continu-
ous beneath the Green River and Bridger formations, and around
the Rock Springs uplift to these widely separated areas.

The more significant of these localities are discussed below :

Rock Springs—West side of Rock Springs uplift, near fork of
Bitter Creek and Little Bitter Creek, about 4 miles southwest of the
city of Rock Springs, collected by R. W. Brown in secs. 7, 8, 17, and
18, T. 18 N., R. 105 W.: Meniscotherium robustum and Coryphodon,
species. The rather fortuitous discovery of these forms in the relatively
thin zone of Wasatchian beds between Rock Springs and Green River
is highly indicative of the upper Knight or La Barge equivalent of
Lost Cabin being represented here between Paleocene and Green
River strata.

Vermilion Creek.—South of Rock Springs uplift on a tributary
to Vermilion Creek (in type Vermilion Creek of King and type Hia-
watha of Nightingale) near line between T. 11 N. and T. 12 N., in
R. ror W., northernmost part of Moffat County, Colo., collected by
writer : Insectivore, possibly Diacodon; Paramys bicuspis; and Hexa-
codus, cf. pelodes. The insectivore found here is not significant.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Paramys bicuspis, according to A. E. Wood, who identified the speci-
men, is suggestive of Lysite; however, Hexacodus, cf. pelodes would
suggest an equivalence with the La Barge fauna.

Black Buttes—East side of Rock Springs uplift, near Black Buttes,
collected in part by Cope and in part by a collector for Marsh:
Coryphodon armatus and Hyracotherium pernix. These give little
information other than a Wasatchian age.

Dad-Baggs.—East side of Washakie basin, exposures along high-
way beneath Tipton tongue, between Dad and Baggs, Wyo., collected
by the writer: Coryphodon, large species, and Hyracotherium, cf.
vasacciense. A small lot of material from near Baggs was collected
by Reeside and Eby in 1924, but this material is not now in the col-
lections. Another collection from near Dad was designated by Wood
et al. (1941, p. 18) as “Dad local fauna” and noted by McGrew (1951,
p. 54) as having been collected by J. LeRoy Kay for the Carnegie
Museum. Of the materials collected by Kay only Hyracotherium was
mentioned. Of the materials which I have observed none are particu-
larly diagnostic; however, I am inclined to regard the horse as of
Knight age rather than lower Wasatchian. It should be noted that
Wood et al. (1941, chart) regard the fauna as late Wasatchian. This
was based on evidence that I have not examined.

Great Divide basin.—About 18 miles west of Rawlins, Wyo., and
about 12 miles northeast of Creston, SEASE4 sec. 32, T. 22 N.,
R. go W., Sweetwater County, collected by E. E. Smith and A. C.
Veatch in 1907 while with the U. S. Geological Survey on the North
Rawlins Coal field: Esthonyx, cf. acutidens; cf. Notharctus venticolus ;
Coryphodon, species; and cf. Heptodon, species. The materials from
this locality in the Great Divide basin are very fragmentary, but cf.
Heptodon suggests Knight, and Esthonyx, ci. acutidens, and cf. No-
tharctus venticolus would appear to place the horizon represented in
the upper Knight or Lost Cabin equivalent. This occurrence is very
near the eastern limit in Wyoming of beds which may be called Knight,
still within the Eocene basin of deposition which includes the Bridger
basin proper.

Cathedral Bluff tongue.—Northwest part of Red Desert, NW. part
of T. 26 N., R. 98 W., collected by R. L. Nace (1939, pp. 17, 26-27)
from 75 feet below top of Cathedral Bluff tongue, and identified by
Simpson: Trogosus or Tillotherium. As Simpson has indicated, the
evidence is rather strong for a Bridger age. However, I am inclined
to regard the Cathedral Bluff and New Fork as equivalent, inasmuch
as so marked a retreat of Lake Gosiute as indicated by these tongues
would effect the entire periphery, unless complicated by tectonic ac-

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 15

tivity involving tilting or folding. In any case there is presumptive
evidence favoring contemporaneity of the occurrences of an extensive
tongue of variegated beds encountered interfingering with the lower
part of the Green River series in various parts of this basin. This
leads to the conclusion that the Cathedral Bluff was deposited during
Lost Cabin time and that Trogosus may well have appeared before the
Bridgerian stage, much as Hyrachyus, and have lived as a contem-
porary of Esthonyx.

GEOLOGIC RELATIONS

Sequence of formations —The sequence of early Tertiary materials
in the southwestern part of Wyoming includes as its lowest unit the
Evanston ; above this the Wasatch group, consisting, as redefined by
Veatch (1907, p. 88), of the Almy, Fowkes, and Knight, is followed
by the Green River and Bridger formations. To the east of the
Bridger basin, around the Rock Springs uplift and along the eastern
margin of the Tertiary basin, as shown by R. W. Brown (1949),
Paleocene beds generally designated as “Fort Union” are exposed,
but whose exact age is not known, having so far produced no known
fossil mammal remains. Washakie beds, overlying Green River in the
Washakie basin, are equivalent to the upper part of the Bridger and
include beds regarded as somewhat later than typical Bridger. To the
south of the Rock Springs uplift, in the basin of Vermilion Creek,
the Vermilion Creek beds of King, as defined by his map (1876, atlas
map 2) are approximately equivalent to the Wasatch of Hayden. The
Vermilion Creek includes Paleocene in the vicinity of the Rock
Springs uplift, and the Tipton and Cathedral Bluff tongues as well as
Knight. The Hiawatha of Nightingale (1930, p. 1023) in the same
general area is in a general way equivalent to the Knight. In the
northern part of the Bridger basin Donovan’s (1950) Fontenelle
tongue of Green River is likely equivalent to the Tipton tongue
farther south, and as noted above, his New Fork tongue of variegated
beds likewise may be Cathedral Bluff tongue, although the identity of
these is yet to be proved. Still farther north in the Hoback basin, beds
which Schultz (1914) mapped as Evanston and undifferentiated
Eocene, and called Hoback formation by the University of Michigan
field parties, have produced a Tiffanian and Wasatchian fauna (Dorr,
1952). The equivalence of these beds with others in the Bridger and
Fossil basin to the west has not been established. The supposition is
that the Paleocene portion may be equivalent to the Evanston, or pos-
sibly to a part of the Almy. The Wasatchian horizon may be lower
Knight, or possibly Fowkes. It is noted, however, that neither Evans-

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

ton nor Fowkes were recognized by Veatch or Schultz elsewhere along
the western margin of the Bridger basin. Should the beds carrying
the Wasatchian fauna in the Hoback basin prove to be a part of the
Knight this would then be the only known place where beds which
appear to be lower Wasatchian in age are included in the formation.

Relations between basins—The geologic relations involved in the
present study are largely those which exist between the sequence of
continental early Tertiary deposits in the upper Green River or
Bridger basins, and those in the small, north-south elongated basin
which includes the type Knight. The latter, which will be referred
to as the Fossil basin, lies in a general way to the west of Oyster Ridge
and includes the high plateau area from the upper reaches of Ham’s
Fork in the north southward beyond the headwaters of Bear River.
On the west it is bounded by the intermittent band of pre-Tertiary
rocks extending in a general way from Tunp Range in the north to
beyond Evanston southward.

Early Tertiary deposition in these two basins appears to have been
entirely separate and independent. As we may see from Veatch’s
(1907) map and sections, the Almy, Knight, and Green River lap
successively onto older rocks along both the eastern and western
margins of the Fossil basin, and along the western margin of the
Bridger or Green River basin, wherever the relation is not actually one
of faulting. And although the pre-Tertiary is much folded and faulted
in the exposed bands along the east and west margins of the Fossil
basin, the Tertiary sediments are for the most part but gently warped
or tilted. However, it is evident that the Laramide activity did not
completely cease by lower Eocene time so that it would be unsafe to
assume that there was entire synchroneity beween the deposits of the
Fossil and Bridger basins. This lack of equivalence is more notice-
able between the Fossil basin and that next toward the west, in the
absence of the Fowkes and in a scant representation of Evanston in
the Fossil sequence. It is probable, therefore, that the Knight to the
east and that to the west of Oyster Ridge are not entirely equivalent
and may have had somewhat earlier or later beginnings and endings,
although I suspect a rather general equivalence.

The same may be said of the Green River series which in the two
basins was deposited in entirely separate lakes. The ultimate reason
for the existence of these two lakes may be the same, or their origins
may be tied to the same structural or orogenic control, but again
their upper or lower limits may not be entirely contemporaneous in
the two basins.

No. 18 LOWER EOCENE KNIGHT FORMATION

GAZIN iL 7

The evidence presented by the fossil mammals in the Knight beds
of the two basins is such as to suggest that the upper limit of the
Knight may have been as much different as the interval between Lysite
and Lost Cabin time. The Lost Cabin fauna in the upper Green
River basin extends from a few tens of feet below the Green River to
100 or more feet, whereas in the Fossil basin so far only a Lysite
fauna has been certainly recognized. In the vicinity of the town of
Fossil, the fauna represented is thought to be equivalent to that of
the Lysite as represented by the type Knight farther south, and was
likewise found 40 to 100 feet below Green River beds. This reason-
ing also leads to supposition that the Green River beds in the Fossil
basin may include somewhat older strata than in the Green River
basin, and that this lake had its origin somewhat earlier in lower
Eocene time.

Should a further and more intensive search of the Fossil basin
demonstrate beyond doubt the absence of beds of Lost Cabin age
immediately underlying the Green River, or interfingering with Green
River to the southward, one might argue the advisability of selecting
a distinct formation name for beds called Knight in the Green River
basin. I would not, however, venture to make such a discrimination,
as the lithology is similar and in all probability there is more or less
overlap if not precise or complete equivalance in time. Continental
deposits are often not clearly definable units, and, of course, notori-
ously unreliable both as to time and lithology when traced laterally
for any considerable distance, even within the same basin of
deposition.

SYSTEMATIC DESCRIPTION OF THE MAMMALIA

MARSUPIALIA
DIDELPHIDAE
PERATHERIUM EDWARDI,? new species
Plate 1, figures 3, 4

Type.—Left ramus of mandible with last two molars, U.S.N.M.
No. 19200.

Horizon and locality—Upper Knight beds, La Barge fauna,
SW4 sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Su-
blette County, Wyo.

2Named for G. E. Lewis, who first called my attention to the particular
locality which has since yielded all the Peratherium material from these beds.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Description.—Size significantly smaller than Peratherium com-
stocki Cope (1884, pl. 25a, fig. 15) from the “Wind River’’ beds,
nearer that of Peratherium marsupium Troxell (1923b) from the
3ridger beds. Cusps of last two molars sharp and well defined, par-
ticularly the entoconid and hypoconulid which in the last molar are
much better defined than in P. marsupium, as represented by National
Museum materials. Length M.-Ms;, 5.5 mm. Transverse diameter of
penultimate molar, 1.8 mim.

A left maxillary portion, U.S.N.M. No. 19206, of a small marsupial
found at the same locality is referred to Peratherium edwardi. The
specimen retains the last two molars, which occlude well with those
of the type. The teeth are rather well worn and show little of diagnos-
tic value. The external margin of the penultimate tooth shows
three small, worn, but about equally well-defined stylar cusps between
the distinct parastyle and metastyle at the angles at the tooth. There
appears, however, but two stylar cusps between the more conspicuous
parastyle and metastyle on the last molar. In Peratherium innomina-
tum Simpson (1928) material in the National Museum collections
from the Bridger, a relatively greater spread between the metastyle
and the preceding stylar cusp is noted in the penultimate and antepen-
ultimate molars. Length of last two upper molars combined, 4.3 mm.
Transverse diameter of penultimate upper molar, 3.1 mm.

PERATHERIUM CHESTERI,® new species
Plate 1, figure 1

Type.—Right ramus of mandible with penultimate molar, U.S.N.M.
No. 19199.

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description—Smaller than hitherto known species of Peratherium
from the Eocene. A little smaller than the Bridger species Peratherium
innominatum Simpson (1928). Penultimate molar length about
1.4 mm, and width about 0.8 mm. The alveoli of the last three molars
measure about 4 mm. Trigonid relatively high and talonid small
with entoconid and hypoconulid much less prominent than in the
larger P. edwardi. The tooth has points of resemblance to an in-
sectivore but the character and position of the hypoconulid and an-
terior cingulum seem certainly marsupial.

3.Named for Chester Gazin, who collected the type specimens of both
P. edwardi and P. chestert.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN I9

INSECTIVORA
LEPTICTIDAE
DIACODON PINEYENSIS,‘ new species
Plate 1, figure 2

Type.—Left ramus of mandible with M, to M, and part of P,,
U.S.N.M. No. 19204.

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description.— Size near Diacodon alticuspis Cope (1875, pp. 11-12)
but with lower molar teeth slightly shorter and distinctly broader.
P, much larger. Length of molar tooth series 8.3 mm., shorter than in
Parictops multicuspis Granger (1910, pp. 250-251) or Diacodon
bicuspis (Cope) (1880a, p. 746; see also Matthew, 1918, p. 575).
Also, lower jaw much shallower than in either. Diacodon pineyensis
is appreciably larger than Diacodon tauri-cinerei Jepsen (1930, pp.
124-126) from the lower Gray Bull beds.

Discussion—A sufficient portion of the heel of Ps; is preserved in
U.S.N.M. No. 19203 to show that the Knight species is Diacodon
rather than Parictops. Moreover, absence of a fourth diminutive cusp
on the heel of the molars is further evidence supporting reference to
Diacodon.

In critically examining the type of Diacodon alticuspis, I find that
the preserved portion of P, is quite unlike this tooth in Diacodon
bicuspis, D. tauri-cinerei, or D. pineyensis. The decidedly short P,
with its narrow, much reduced talonid in D. alticuspis might justify
resurrecting the generic name Palaeictops for D. bicuspis, etc. Mat-
thew’s reasons for first erecting ® and then dropping the name Pa-
laeictops are not given. However, the differences here noted are as
great as, if not greater than, those between Parictops multicuspis and
Diacodon bicuspis. I (1949, p. 221) earlier regarded Parictops as
probably valid, but the significance of its characters may be no greater
than those which seem indicated for Palaeictops. It should be noted,
moreover, that in the character of P; Diacodon tauri-cinerei makes a
distinct approach toward Parictops multicuspis. It is anticipated that
new collections currently being made by Dr. Simpson from the San
Jose formation in New Mexico will reveal additional material of
Diacodon alticuspis so that this form will be more clearly defined and
the genus better understood.

4Named for the town of Big Piney, Wyo.
5 Granger (1910, p. 250) has incorrectly attributed this genus to Cope.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

DIACODON, cf. ALTICUSPIS Cope

There is in Princeton’s collection, from about 100 feet below Green
River beds, 3 miles northwest of Fossil, Wyo., a lower jaw portion,
P.U. No. 16171, representing a species of Diacodon very close to D.
alticuspis Cope. The species is clearly not Diacodon pineyensis. Com-
pared to the latter the cheek teeth are narrower, particularly the
talonids, and the paraconid is placed lower with respect to the much
elevated protoconid and metaconid. The narrowness of the talonid is
most emphasized in P, in which the entoconid is much subdued and
the basin restricted. The teeth, except for P,, are slightly larger than
in D. tauri-cinerei and the trigonids of the molars are higher, except
for the paraconid, and broader. Comparison with D. alticuspis sug-
gests a close relationship, but one which is inconclusive owing to the
fragmentary nature of the type.

MIXODECTIDAE
CYNODONTOMYS KNIGHTENSIS,® new species

Plate 2, figure I

Type.—Leit ramus of mandible with Py-M;, U.S.N.M. No. 19314.

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description—Size close to that of Cynodontomys latidens Cope
(1882a, pp. 151-152), much smaller than more nearly contemporane-
ous Cynodontomys scottianus (Cope) (1881a, pp. 188-189). Ps
with paraconid distinct and low, and with metaconid almost as large
as protoconid and well separated from it though not so much so as in
Microsyops. Talonid of P, broad and fully molariform, though
slightly narrower than in molars, with protoconid, entoconid, and
hypoconulid sharp and well defined. P, distinctly more progressive
than illustrated (Matthew, 1915c, figs. 45, 46) for Cynodontomys
latidens Cope.

Discussion.—In all there are 13 specimens of this form, all but
one are lower jaws and, except for two, are from the same exposures
as the type. One lower jaw and an isolated upper molar were obtained
southeast of Big Piney, on the east side of the Green River. This
interesting species, except for its smaller size, is as fully developed as
Cynodontomys scottianus in the progressiveness of its fourth lower
premolar, as might be expected in this horizon. Cynodontomys scot-

6 Named for the Knight formation.

no. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 21

tianus, however, is not represented in this fauna, nor is the relatively
gigantic Cynodontomys lundeliusi White (1952, pp. 191-192) from
the Lost Cabin beds in the Boysen Reservoir area. Though C. knight-
ensis and C. scottianus are from distinctly separate basins of de-
position during upper Wasatchian time, the geographic distance be-
tween the occurrences is not particularly great. It seems probable
that the difference in species is to be attributed to ecologic differences.

MEASUREMENTS OF TYPE LOWER JAW OF
Cynodontomys knightensis, NEW SPECIES, U.S.N.M. NO. 19314

mm.
EA SPUSCNEENS,) ip LO av isa INC ara oiie crvevatcotie are Oe neratern aon taltoua ataxic a eG 20.5
PEP e AST Me aah oe feccrc a: cialis ata Wa stare, « CR #ccesc ores olin GY aE 15.2
M, to Ms, ATT Meo) nabs oe ach Sas Sea er S65 Soe. aot oid -ancin OCIS tC CRITE 11.5
Pept or yaw beneath Me Caternally).......6.swccs ac cee caccnte 1

CYNODONTOMYS, species

A portion of a left lower jaw including M, and Ms, in the collections
of the American Museum, from the bluffs of the Bear River south of
Knight station is identified as Cynodontomys. M, in this specimen has
the same proportions as in C. knightensis, but Ms is relatively a little
shorter as in Cynodontomys angustidens. This specimen, A.M. No.
12836, has been labeled Cynodontomys latidens? but in the absence
of P, the progressiveness of the dentition cannot be clearly deter-
mined. An unusual feature of the specimen is the remarkable flange-
like development of the outward- and forward-turned anterolateral
margin of the ascending ramus. An approach to this condition is seen
in the prominence of this crest in certain of the specimens of C.
knightensis.

TILLODONTIA
ESTHONYCHIDAE
ESTHONYX, cf. ACUTIDENS Cope

Two incomplete, isolated upper molars from variegated beds be-
tween Big Piney and La Barge, Wyo., and a single lower molar from
12 miles North of Big Piney, may well represent the species Esthonyx
acutidens Cope (1881a, pp. 185-186). The lower tooth is complete
and little smaller than in the type of E. acutidens. There is much
variation in size of Esthonyx material from the Lost Cabin beds but
the average is distinctly larger than the older E. bisulcatus; more-
over, the posterior wall of the trigonid of the lower molars tends to
be somewhat less oblique and the talonid basin a trifle larger in
E. acutidens.

i)
iS)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.) a7,

In passing it should be noted that the type of the species, Esthony«
spatularius, erroneously recorded as coming from the Wind River
beds, is beyond reasonable doubt from the Gray Bull beds of the Big
Horn basin. The history and explanation of this error are covered in
manuscript now in preparation on the tillodonts, and briefly stated in
this paper under the section on Didymiciis.

ESTHONYX, species

The outer portion of an upper molar in the Knight station material
in the American Museum is recognized as belonging to Esthonyx.
The specimen is too incomplete to warrant specific comparison. An
upper molar tooth in the collection from the New Fork tongue also
represents Esthonyx. The latter tooth is short anteroposteriorly and
relatively wide transversely. I am uncertain as to whether or not it is
Esthonyx acutidens.

PRIMATES
ADAPIDAE
PELYCODUS>? species

A fragment of an upper molar, including only the outer half, in the
American Museum collection from the Knight beds near Knight sta-
tion in southwestern Wyoming, is only slightly larger than M? in
Notharctus limosus described below, but has the mesostyle no better
developed than in Pelycodus, hence is referred tentatively to the latter
genus,

NOTHARCTUS LIMOSUS,’ new species

Plate 1, figures 6-8

Type.—Portions of both rami of the mandible with left P:-P,, and
right M,-Msg, inclusive, U.S.N.M. No. 19294.

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 117 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description.—Size of molars a little less than in Notharctus
nunienus (Cope) (i88ia, p. 187) with premolars much smaller and
depth of jaw less, particularly beneath premolars. P, with metastylid
well developed and broadly spaced from protoconid. Upper cheek
teeth anteroposteriorly short, most noticeably in the medial and
lingual portions. Molars relatively broad transversely with distinct
mesostyle and hypocone on M? and M?.

*From Latin limosus, muddy, with reference to Muddy Creek where the
type was found.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 23

Discussion.—The smaller of the notharctine primates in the La
Barge fauna is represented by about 24 specimens, the majority of
which were obtained from exposures about 12 miles north of Big
Piney.

The lower molars average noticeably smaller in size than those in
American Museum specimens of Notharctus nunienus from the Lost
Cabin beds, and in only two specimens are teeth found as large as in
the smallest of NV. nunienus. The length of Ms, for example, has a
range of from 4.9 to 5.3 mm. in N. nunienus, and in N. limosus the
range would be 4.3 to 4.7 mm. except that one specimen measures 5.0.
The average in the two species is 5.2 and 4.6 mm., respectively. The
molar teeth are otherwise not particularly distinctive. The paraconid
is strong on M,, but highly variable on M, and M;. Ms has two cusps
in the position of the hypoconulid much as in N. nunienus, although
this has been noted also in Pelycodus.

The difference in size of premolars is more noticeable, and P; in
particular is relatively much smaller than in N. nunienus. Pa,
though of distinctly smaller size than in N. nunienus, has a metaconid
that appears to be relatively more widely separated from the proto-
conid and exhibits a conspicuous, sharply conical hypoconid. This
cusp is also distinct on P3. In N. nunienus material the accessory cusp
or hypoconid on P; and Py, is generally weak or blunt. In one speci-
men of N. nunienus, A.M. No. 4736, a rather distinct entoconid was
observed on P,. A rudiment of such a cusp was noted in U.S.N.M.
No. 19290 of N. limosus.

A well-preserved maxillary portion, U.S.N.M. No. 19293, of
Notharctus limosus, includes P? to M’. The upper teeth are notice-
ably less in their fore and aft dimension, taken through their middle
and lingual portions, in comparison with A.M. No. 4735 of N.
nunienus, giving them a less rounded appearance, but they are nearly
as wide transversely. As in N. nunienus, the mesostyle is distinct on
M? and M2, and somewhat weaker on M*. The hypocone is as well
developed on M? and M?, but obscured through wear on M®.

MEASUREMENTS OF TYPE LOWER JAW OF
Notharctus limosus, NEW SPECIES, U.S.N.M. NO. 19204

mm,
WPM po tdAC Lasoo, Ak Arvin ats pod occ pehetoua= apie sus sielteGe = oa) erate ase jn.seraie 14.4
P;,® anteroposterior diameter: greatest transverse diameter.... 2.9: 2.0
Ps & i s #3 dere ALO a7)
M;, f a = . oes a
Mz, a if. iS $0) Siw AARSS
Mz, " > 4 e Ys eevee eR aS

8 The type specimen is at the upper limit for the sample in size of premolars.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

NOTHARCTUS VENTICOLUS Osborn

The larger of the notharctine primates in the La Barge fauna is
represented by a maxilla, U.S.N.M. No. 19288, with P*-M® and five
lower jaws. The size of the teeth in these specimens corresponds well
with those of Notharctus venticolus Osborn (1902, p. 195). In the
upper molars the mesostyle is prominently displayed, but the hypo-
cone, though apparently well developed on M’, appears subdued on
M?. It is evident, however, that somewhat greater wear would pro-
duce a large distinct lake in this position. The lingual portions of all
the upper teeth preserved in this specimen appear relatively broader
anteroposteriorly than in the N. limosus maxilla. P* and M® are dis-
tinctly less nearly triangular.

ANAPTOMORPHIDAE
ABSAROKIUS NOCTIVAGUS Matthew

This species is apparently represented in the La Barge fauna
materials by three lower jaw portions, one of which, U.S.N.M. No.
19198, exhibits Dp, with what appears to be Py, as well as P, and Ms,
just erupting. The first two molars are in position but unworn. If one
may judge by the positions of the two erupting premolars, P; is yet
buried beneath the long slender roots of Dps. Two small alveoli
anterior to the erupting P.? are presumably for incisors, or incisor
and canine. There is no evidence for an enlarged front tooth as in
Tetonius. Matthew (1915c, pp. 463-465) was apparently correct in
his interpretation of the condition in Absarokius abbotti (Loomis)
and fully justified in distinguishing Absarokius.

The teeth in No. 19198 compare very closely to those in the type of
A. noctivagus but are very slightly narrower. In U.S.N.M. No.
19196, M; is a little smaller than in this type, about the length but not
so slender as in a specimen of Absarokius abbotti in the American
Museum collections. A third specimen in the collections from north
of Big Piney, U.S.N.M. No. 19197, includes an M; a little larger
than in the type of A. noctivagus, and the paraconid of this is much
less lingual in position. It is possible that the latter specimen does not
represent this genus.

PARATETONIUS? SUBLETTENSIS,® new species
Plate 1, figure 5

Type.—tLeft ramus of mandible with P,-M,, inclusive, preserved,
U.S.N.M. No. 19205.

® Named for Sublette County, Wyo.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 25

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description.—Size smaller than Paratetonius? tenuiculus (Jepsen)
(1930, p. 126), much smaller than Paratetonius musculus (Matthew)
(1915¢, p. 463) or Paratetonius steini Seton (1940) from the older
Gray Bull or Lysite horizons. P, not enlarged but exhibiting a well-
defined paraconid, a weak but distinct metaconid, a short but broadly
basined talonid with but a very weak posterointernal cusp or entoconid
and a somewhat better-defined posteroexternal cusp or hypoconid.
Molar cusps more marginally placed than in Tetonius, as observed by
Seton for Paratetonius. Paraconids lingual in position, and in M,
clearly defined and close to the metaconid. As shown by alveoli, P;
is two-rooted, not fused as in P. musculus, and P, apparently has but
a single large root, relatively larger than in P. musculus. Anterior
to the root of P, is a very small but deep alveolus, probably for P,,
but possibly for an anterior root of P2, posteroexternal to the relatively
large alveolus presumed to be that of the canine.

Discussion.—As anticipated by Matthew in his description of
Tetonius musculus, Paratetonius Seton appears to be a clearly dis-
tinct group, less specialized than Tetonius, and including forms in all
horizons of the Wasatchian stage. The latest of these, Paratetonius?
sublettensis, if correctly referred, is the smallest and in some respects
may be the least specialized.

In addition to the probable dental formula, the moderate-sized,
relatively low-crowned P, implies less progressiveness than, for ex-
ample, Tetonius homunculus. On the other hand, the form of P, is
distinctly more progressive in its approach to that of the molars, a
quite different trend.

It is interesting to note that Loomis’s form, Anaptomorphus mini-
mus (1906, p. 278), from Gray Bull beds of the Big Horn basin,
appears scarcely to have been mentioned since its description, except
for having been placed questionably in synonymy with Nyctitheritum
celatum by Matthew (1918, p. 604). The illustration is not too clear,
so that generic reference cannot be verified but in size it may be
smaller than Paratetonius musculus. The dimensions given by Loomis
are slightly less than for lower teeth in a Gray Bull specimen in the
National Museum collections, No. 19154, tentatively referred to
Paratetonius? tenuiculus, but close to those of Paratetonius? sub-

lettensis.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117,

MEASUREMENTS OF TYPE LOWER JAW OF
Paratetomus? sublettensis, NEW SPECIES, U.S.N.M. NO. 19205

inm
Thensthy WPsitom Mas beh. ey litha a tiee etka avers ey aie ey A pe een 4.5
Length: My tos Ma scimele: ici. t outs seeds om eee ese ee eetea iene ee
WV fdas alias ac acetate losers ces sasstct acaea ke nancy sae iver teen rt aN ERC eg cs Tipe.
Depth Of jaw beneath Me intertally....+-s oc eee eee ee cae 3.1
TAENIODONTA
STYLINODONTIDAE

Undetermined stylinodont
Plate 2, figures 2-5; plates 3, 4

The articulated portions of two hind limbs and a fore limb, with
feet, and including some vertebrae from exposures of upper Knight
12 miles north of Big Piney, are recognized as representing a taenio-
dont. No skull or jaw material, except for some finely broken and
scattered fragments of teeth, was encountered. So far the only deter-
minable taeniodont material from this horizon is the Lost Cabin
Stylinodon? cylindrifer (Cope) (1881a, pp. 184-185) from the Wind
River basin. It is reasonable to suppose that the La Barge fauna in-
cludes that species but this cannot be established on the material at
hand.

The finding of this material is indeed fortunate as it gives us in-
formation previously not known on the structure of the hind foot in
this family, and gives a certain amount of skeletal data none of which
had been recorded before from the Lost Cabin stage.

Fore limb.—The limb bones are rather badly crushed and those of
the fore limb very incomplete. The greater part of an ulna is pre-
served and though incomplete distally, so that its length cannot be
determined, it resembles rather noticeably the ulna figured by Marsh
(1897, fig. 5) for Stylinodon muirus.

Fore foot—The fore foot is represented by the scaphoid, pisiform,
unciform, the fourth and fifth metacarpals and the distal part of the
third, together with the phalanges of the second, third, and fourth
digits. The scaphoid is a relatively small triangular wedge with the
large proximal or superior facet for the radius, converging forward
and medially with the somewhat smaller distal facet. These are sepa-
rated laterally by a still smaller lenticular facet for the lunar. The
angle between the lunar and distal facets is about 90°, but between
the lunar and radial facets is much more acute, more as anticipated by
Matthew (1937, fig. 66) for Psittacotherium than as interpreted in

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 27

rough outline by Patterson (1949, p. 253, fig. 4c) for Stylinodon
mirus. It cannot be determined from the material at hand whether
the moderate-sized “pear-shaped distal facet articulated with the
trapezium, or, as Patterson suggests, the trapezoid. This bone, how-
ever, so very closely resembles the scaphoid in Trogosus, the distal
facet of which articulates with the trapezium, that a reasonable doubt
persists. ;

The pisiform is deeply expanded dorsoventrally in its posterior
portion and anteriorly exhibits a large, concave articular surface for
the ulna and ventrally a much smaller facet for the cuneiform. The
anterior extremity is expanded medially although this projection is
not complete.

The unciform is a relatively large carpal bone which articulates
distally with the entire proximal end of the fourth metacarpal. It
is incomplete laterally so that the surface for the fifth metacarpal is
missing. There is no certain evidence that the unciform was in contact
with the third metacarpal. The proximodistally long facet on the
medial side of the unciform seems to have articulated only with the
magnum as Patterson has shown. Proximally, incomplete facets for
the lunar and cuneiform, at a sharp angle to one another, are pre-
served.

The fourth metacarpal, about 35 mm. in length, is a sturdy bone
with broadly expanded extremities, but with a constricted shaft. The
articulating surface for the fifth metacarpal is rather broadly arcuate
and undercut distally in a somewhat conventional way, but facing
distinctly more palmar than in many unguiculate types. The proxi-
mal surface is nearly triangular and the broad and deep distal articular
surface for the proximal phalanx is evenly rounded dorsoventrally
but with almost no evidence of a keel. The 2d and 3d metacarpals are
seen to have saddle-shaped distal extremities.

The fifth metacarpal is about two-thirds the length of the fourth
and has a moderately robust proximal portion with broad articula-
tion with the fourth metacarpal. A facet is seen for articulation with
the unciform but this surface is incomplete in the specimen at hand.
The distal portion of the fifth metacarpal is very much reduced with
but a very small articular surface for the first phalanx, about half the
width of that in the fourth.

The phalanges of the second to fourth digits are broad and deep
but much shortened proximodistally. Only the proximal portions of
the clawed distal phalanges are present, but these are transversely
narrow and extremely deep. The largest of the claws is approximately

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

18 mm. across the widest part of the base proximally, but is about
38.5 mm. deep.

Hind limb.—The stylinodont hind limb and foot, previously un-
known but for the meager foot remains belonging to Psittacotherium
and very fragmentary limb material of Ectoganus, are represented by
both right and left sides in the Knight material. In the left limb all
elements are included, but the tibial and tarsal portions are rather
poorly preserved. The right side is in a much better state of preserva-
tion; however, the femur, cuboid, part of the external cuneiform, and
the proximal portions of the second and fifth metatarsals are missing.
Except for the proximal articulating surface on the second metatarsal
and much of the cuboid, these can be interpreted from the left side.

The femur, approximately 220 mm. long as measured from about
the crest of the greater trochanter, is crushed and distorted, so that
detailed description is not feasible. The head is lacking but the greater
trochanter is well developed. It is broadly expanded anteroposteriorly
and its roughened external surface extends over 80 mm. distally.
There does not appear to be a third trochanter but the lesser trochanter
beneath the head, though broken off, was evidently well developed
for possibly as much as 50 mm. along the shaft. The distal or condylar
portion of the femur though transversely crushed would appear to be
narrow and moderately deep anteroposteriorly.

The proximal portion of the tibia likewise appears somewhat
crushed transversely. Nevertheless, the proximal extremity may be
relatively deep anteroposteriorly and transversely narrow, particularly
just below the articular portion. The external articular surface is
relatively small and immediately beneath its posterior extremity and
facing distally is a facet for the fibula. The inner surface of the
proximal portion of the tibia is rather flattened and the internal
condyle is much more produced anteroposteriorly than the outer. A
prominent ridge extends downward on the posterior margin of the
shaft from the external condyle, terminating distally in a prominent
and unusual-appearing process *° about 36 mm. below the articular
surface. The anterior margin or cnemial crest slopes more or less
evenly toward the distal portion of the shaft with no clearly defined

10 This process may well have accommodated in part more than one im-
portant muscle. Its direction suggests the soleus, participating in the Achilles
tendon. However, the groove posterior to the internal malleolus is well developed
for the tibialis posticus or flexor longus digitorum, or both, which also may
have originated in part on this process. The flexor muscles were obviously
strong, to judge by the claws, and a well-developed tibialis posticus might account
for a similarly unusual development of the posterointernal margin of the navicu-

lar as a point of insertion. The latter muscle would serve as an extensor of the
tarsus and in supination of the foot.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 29

lower limit on the material at hand, but the proximal tuberosity stands
out ruggedly and well in front of the condylar surfaces. The distal
portion of the tibia appears normal and undistorted. The internal
malleolus is very well developed and extends downward to cover a
large segment of the inner wall of the astragalus. Also, the inner
condylar surface of the distal end of the tibia projects much farther
posteriorly than the outer.

The fibula is a slender shafted bone with enlarged extremities.
Also, the shaft is somewhat curved, longitudinally, with the concavity
forward so that the distal extremity or external malleolus is well
forward and actually ahead of the tibia. The external malleolus is
transversely broad anteriorly and on its inner side exhibits a large
and elongate surface facing inward, ventrally, and somewhat pos-
teriorly for articulation with the forward, proximal portion of the
external wall of the body of the astragalus.

Hind foot.—The hind foot for the most part has the general appear-
ance of a primitive unguiculate type, but with a combination of dis-
tinctive characteristics evidently more or less peculiar to the Taenio-
donta. The more notable peculiarities include a transversely, much
recurved arrangement of the tarsus and digits. The cuneiforms and
proximal extremities of the metatarsals present a triangular aspect so
that articulation between them from side to side tends to bring the
first and fifth digits relatively close together in a plantar direction.
With this arrangement the animal’s metatarsals were evidently main-
tained nearly upright in standing, with the weight to the ground
carried largely by the distal extremities of the metatarsals through
their large sesamoids, and only in part by the proximal phalanges. It
is also interesting to note that the first and fifth digits, though a little
shorter than the others, are relatively unreduced, as compared with
those of the front feet, and although the various digits diverge notice-
ably from one another, the first appears to diverge from the second
somewhat more sharply. The characteristics of the individual ele-
ments of the foot are discussed in more detail below.

The astragalus in proximal aspect rather resembles that in the
larger cats, but with the trochlear surface distinctly raised, not con-
tinuing more or less smoothly into that of the neck of the astragalus.
It is not far from the conception outlined by Matthew (1937, fig. 67)
for Psittacotherium, although with less anteroposterior extent of the
trochlea for the tibia. In plantar aspect, the astragalus is somewhat
more distinctive. The large ectal or peroneal facet is less concave than
in cats or bears. Posteriorly it converges rapidly with the trochlea
rather than extending parallel to it, giving the external wall of the

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

body of the astragalus a triangular rather than lunar outline. An-
teriorly the ectal facet turns sharply upward for a short distance.
The sustentacular facet is much smaller, slightly concave antero-
posteriorly, and elongate, extending from near the navicular facet
medially, posteriorly, and plantad with respect to the ectal facet. The
head of the astragalus articulates broadly with the navicular and
externally in a small part with the cuboid.

The calcaneum is apparently the most distinctive bone in the pes.
Its most noticeable characteristic is the dorsoventral expansion of its
tuberosity or tuber calcis, recalling on a small scale somewhat the
appearance of the posterior margin of this element in certain of
the edentates, such as Hapalops. Ventrally the expansion projects
forward, presumably in part for attachment of the flexor brevis digi-
torum which would be inserted into the second phalanges. Anteriorly
the calcaneum exhibits on its superior margin the large ectal facet for
the astragalus. Anterior to this and meeting the upturned anterior
margin of the ectal facet at a sharp angle is a transversely elongate
and concave facet facing forward and medially for the cuboid.
The outer surface of the calcaneum is developed anteriorly into
a prominent forward projection or process which carries the forward
portion of the cuboid facet on its inner surface. The sustentacular
portion of the calcaneum is missing.

The navicular is distinguished by its elongation, extending inward
well beyond its articulation with the head of the astragalus and the
internal cuneiform. The knoblike ventrointernal prominence on the
navicular may well represent a sesamoid commonly found in certain
groups of mammals on the tibial side of the tarsus, which has here
become co-ossified with the navicular. The proximal surface of the
navicular is broadly concave with a ventrointernally elongate facet
for the head of the astragalus. The external facet for the cuboid is
about at right angles to the surfaces for the astragalus and external
cuneiform, and moderately short dorsoventrally. Its direction is
distinctly acute to that of the dorsal surface of the navicular. Dis-
tally, the three facets for the cuneiforms occupy somewhat less than
two-thirds the long diameter of the navicular. These facets are of
approximately equal width dorsally, but that for the internal cunei-
form is the more convex dorsoventrally and turns sharply proximal
internally.

Little can be determined of the characteristics of the very poorly
preserved cuboid in the left foot; however, from the articular facets
of the adjacent bones of the right foot it was clearly of good size.
Proximally, it articulated broadly with the calcaneum and extended

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 31

onto the externoplantar margin of the head of the astragalus. In-
ternally it made contact with the nearly erect and coplanar surfaces
of the external walls of the navicular and external cuneiform. Dis-
tally, it was evidently broad and covered much of the proximal
extremity of the fourth metatarsal and apparently a somewhat larger
surface on the fifth metatarsal.

The middle and external cuneiforms are of about equal size and
present a nearly rectangular appearance in dorsal view. Their
proximal and distal aspects are more nearly triangular, the external
cuneiform more so than the middle cuneiform. On their distal surfaces
each laps slightly over the next metatarsal external to that directly
distal to it. The internal cuneiform is not well preserved but is dis-
tinctly larger than the other two. Proximally its articular surface for
the navicular is markedly concave and externally it makes contact
only with the proximal portion of the inner wall of the middle cunei-
form. Distally its articular surface is broad, deep, and slightly convex
for the first metatarsal.

The metatarsals are a little shorter, with shafts somewhat con-
stricted, and extremities expanded in comparison with the partial
hind foot of Psitiacotherium figured by Matthew. The first meta-
tarsal is the shortest, about five-sevenths the length of the third
metatarsal, but is more robust and the proximal extremity is some-
what larger.. The first metatarsal apparently does not articulate with
the second. The latter is intermediate in length between first and
third metatarsals and has a somewhat broader and flatter shaft than
the third. The distal extremity is also a little wider but more com-
pressed toward the first. The third and fourth metatarsals are about
the same length, 41 mm., but the fourth metatarsal articulates in a
slightly higher position in the foot than the third. The proximal
extremities of both are distinctly triangular with the broadest base
forward. The shaft and distal extremity of the fourth is slightly
wider and shallower than the third. The fifth metatarsal is a little
longer than the first, but much less robust, and highly distinctive in
appearance. The proximal portion has a wide external flare and the
dorsoventrally much-compressed shaft exhibits a much more broadly
concave plantar surface than the other metatarsals. The proximal
extremity articulates with the full depth dorsoventrally of the external
surface of the fourth metatarsal and has a relatively large articular
surface for the cuboid. Distally the articulation with the first phalanx
is transversely wide but not deep. All the metatarsals show well-
rounded distal articulations and sturdy sesamoids, and only the
slightest suggestion of a median keel.

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL) LE?

The intermediate phalanges of all the digits of the hind foot are
much shortened, more so than in Psitiacotherium, and the proximal
of these in digits 2 to 5 are scarcely more than wedges with the thin
edge down. Those adjacent to the clawed phalanges show evenly
rounded, saddle-shaped articular surfaces distally. The claws are
much more robust than in Psittacotherium, but not nearly so deep and
relatively less compressed transversely than in the fore foot of
the Knight individual. Those in the first and fifth digits are fully
developed.

EDENTATA
EPOICOTHERIIDAE?
PENTAPASSALUS,!! new genus

4 ?
Generic characters——Dental formula: I--, CL, P*, M-3.
i I

Teeth much as in Tubulodon, but last premolar and M, possessing a
single, conical root. M, alone of the lower series two-rooted. Cranial
portion of skull relatively short and broad, more as in Epotcotherium
than as in Palaeanodon or Metacheiromys. Bullae ossified, well in-
flated and posterior in position. Palate not extended posteriorly and
pterygoids not continuous with bullae.

Type species.—Pentapassalus pearcet, new species.

PENTAPASSALUS PEARCEI,!2 new species
Text figures 2-4

Type.—Large part of skeleton, including skull and lower jaws,
U.S.N.M. No. 20028.

Horizon and locality—Upper Knight beds, La Barge fauna, east
side of Green River, southeast of Big Piney, sec. 14, T. 29 N., R. 111
W., Sublette County, Wyo.

Specific characters——Size near Tubulodon taylori, but with teeth
a little smaller and jaw somewhat deeper. Skeletal portions correspond
closely in size to those of Metacheiromys tatusia. Other specific char-
acters not distinguished from those of the genus.

Material—The type of Pentapassalus pearcei is a somewhat crushed
but remarkably complete skull and lower jaws, together with both
humeri, portions of both radii, an ulna, femora, parts of the tibiae,

11 From Greek 7évre, five, and wéocados, a peg; with reference to the five
peglike postcanine teeth in the skull.
12 Named for Franklin L. Pearce, who found the type specimen.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 33

incomplete scapula and pelvis, a few vertebrae, ribs, and foot bones.
In addition to the foregoing, there is in the collection a left lower
jaw portion, U.S.N.M. No. 19214, with the last molar and the alveolus
for the molar immediately preceding, which was found about 12 miles
north of Big Piney, Wyo.

Description of skull—Among living forms the skull of Penta-
passalus pearcei may best be compared with that of the small Argentine

ma
as
i .
\.

Fic. 2.—Pentapassalus pearcei, new genus and species: Skull and mandible
(U.S.N.M. No. 20028), type specimen, dorsal, posterior, lateral, and ventral
views, natural size. Upper Knight, lower Eocene, Wyoming.

armadillo, Zaédyus pichiy Desmarest, rather than with the peculiar
Chlamyphorus truncatus or Burmeisteria retrusa. In superior view
(fig. 2) the cranial portion of the skull of P. pearcei is wider, and the
rostrum much shorter and less attenuated anteriorly than 2. pichty.
The anterior extremity of the rostrum is damaged but its extent can
be determined by the nasals and lower jaws. Posteriorly the rostrum

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II7

widens abruptly across the frontals, but apparently not to the extent
in Z. pichiy. The sutures of the fossil skull are for the most part well
closed so that no great amount of information on the limitations of
the individual bones can be ascertained. However, the indistinct
suture between the premaxilla and maxilla appears to curve backward
superior to the bulge of the canine. Also, the frontoparietal suture
extends almost directly across the cranium at the orbital constriction.
There is no sagittal crest, but the weak superior margins of the
temporal fossae approach closely and extend nearly parallel fore and
aft, somewhat as in Zaédyus. The presence or absence of an inter-
parietal cannot be determined.

In lateral view (fig. 2), the skull of P. pearcei exhibits moderate
depth posteriorly, relatively a little less than Zaédyus pichiy, but
becomes noticeably shallow immediately anterior to the bullae. The
midportion of the skull is somewhat crushed, but the basicranium
makes a decidedly more acute angle with the palate than in the arma-
dillo or Orycteropus. In the latter forms these surfaces are more
nearly parallel. The much shorter rostrum in P. pearcei does not taper
anteriorly so rapidly. The lachrymal portion of the skull is rugged
and participates in a marked lateral widening of the skull at the
anterior margin of the orbit, much as in Zaédyus, and the lachrymal
foramen is conspicuous on the crest of the zygoma immediately be-
neath a rugose prominence of the lachrymal bone. This foramen is
more forward in Orycteropus and some material of Dasypus. The
anterior opening of the infraorbital foramen is large, single, and
immediately above M'. The zygoma is slender with the squamosal
portion overlapping the jugal much as in Orycteropus. The contact
of these elements is quite unlike that in Dasypus, and the zygoma is
much weaker than in Zaédyus.

In posterior view (fig. 2) the skull is seen to be broad and shallow,
actually resembling more closely the much larger Orycteropus than
any of the armadillos. The resemblance is further emphasized by
the way in which the somewhat more inflated mastoid portion is dis-
played in this view, and in the extension of the well-developed
lambdoidal crests outward, above, and lateral to the mastoid promi-
nence. The foramen magnum in P. pearcei is notched dorsally some-
what as in E poicotherium.

The ventral aspect (fig. 2) of the P. pearcei skull is highly inter-
esting and shows in its anterior part a short, broad palate in contrast
to that in armadillos and aardvark. Although somewhat destroyed
posteriorly, the palate, as in Zaédyus, apparently did not extend much,
if any, posterior to the tooth row. The palatal margin of the posterior
narial opening was concave and there is a good indication that it was

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 35

also curved downward, somewhat as in Orycteropus. The pterygoids
are partially damaged but apparently turn outward ventrally rather
than inward, possibly more so than in Zaédyus or Orycteropus, but
quite unlike Dasypus. The palatopterygoid crests were much longer
than in Zaédyus, more as in Orycteropus but relatively not so widely
separated. These crests also were not continuous with the bullae as
in Burmetsieria or Zaédyus.

The foramina in the alisphenoid of Pentapassalus may not be dis-
tinctive, but the identity of all has not been clearly determined. The
foramen ovale is in a normal position just above the posterior extent
of a subdued lateral sweeping branch of the pterygoid crest. Im-
mediately anterior to the foramen ovale and within the same general
depression is a small foramen, noted also by Simpson (1927, p. 289)
in Epoicotherium, which may be the posterior opening of an alisphe-
noid canal but is in all probability a venous canal, which extends trans-
versely through the basisphenoid in both armadillos and Orycteropus.
The presence or absence of an alisphenoid canal cannot be determined
in Pentapassalus without damage to the skull. It is absent in arma-
dillos and Orycteropus, although Gregory (1910, p. 335), following
Weber (1904, p. 414), denoted its presence in the latter—a mistaken
identification of the foramen ovale, which was corrected in a later
edition of Weber’s work. Just ahead of the small venous foramen
noted above is a circular foramen, somewhat smaller than the foramen
ovale which may be a foramen rotundum; however, the position of the
alisphenoid-orbitosphenoid suture has not been determined with cer-
tainty so that this may well be the sphenoidal fissure. Immediately
forward of the latter opening, the bone shows an elongate, natural-
appearing margin directed anterodorsally, such as might be expected
of the posterior and outer lip of a sphenoidal fissure, but from there
forward the orbital cavity is damaged so that the orbital foramen can-
not be seen.

The broad basicranium is characterized by the markedly inflated
bullae, the medial walls of which are more nearly parallel than in
Zaédyus. The anteromedial margin of the bulla is deeply grooved
from the medial wall forward to the foramen lacerum medium for the
internal carotid. This groove is not covered medially, nor is the bulla
perforated ventral and lateral to this position as in Zaédyus. This
perforation may well be the eustachian canal in the latter. In
Pentapassalus the eustachian foramen may be nearly confluent with
the foramen lacerum medium ventrally, or be represented by a small
opening at the anterior margin of the bulla well separated lateraily
from the foramen lacerum medium. The external auditory meatus is
large and opens immediately beneath the lower extremity of the lamb-

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117

doidal crest. The glenoid surface for articulation with the lower jaw
is well forward from the external auditory meatus and terminates
posteriorly in a much-subdued postglenoid process, which carries on
its posterior surface a normal postglenoid foramen. In Dasypus there
is no postglenoid process and the articular surface terminates pos-
teriorly at the opening for the ear. In Zaédyus and Orycteropus the
postglenoid process of the squamosal is a thin sheet of bone which
also forms the anterosuperior margin of the external auditory meatus.
The bulla in Dasypus and Orycteropus is not ossified, only an annular
ring is present, whereas in Zaédyus the bulla is ossified, but unlike
Pentapassalus does not participate in the margin of the external
auditory meatus as this is formed entirely by the mastoid and squa-
mosal. Posterolateral to bulla the margin of the skull is formed by
the well-inflated mastoids, unlike the armadillos but somewhat more
as in Orycteropus. Posterior to the bulla the foramen lacerum pos-
terius is evident, and the condylar or hypoglossal foramen is distinct
and posterior to it. The stylomastoid foramen would be distinctly
marginal in position, immediately posterior to the external auditory
meatus. Its exact position and that of the hyoid articulation are some-
what indistinct in the fossil as a result of minor fracturing. The
occipital condyles are not so widely separated as in armadillos and
these do not so nearly encroach on the mastoids, or bullae.

Comparison of the skull of Pentapassalus pearcet with other fossil
types shows a marked resemblance to various previously described
palaeanodonts; however, important differences were noted which
clearly distinguish the genus. The cranium of P. pearcei is almost
as wide as in Gray Bull Palaeanodon ignavus Matthew (1918, figs.
39, 40) but shorter, and the total skull length is very much less. In
keeping with the relatively shorter cranium the bullae are placed
farther back than are the incompletely ossified bullae or annular rings
in P. ignavus. The rostrum in P. ignavus is markedly elongate but
not attenuated as in Dasypus. The dentition of P. ignavus is much
farther forward than in P. pearcei and the number of teeth appears
to have been less. In an occipital view the mastoid portion is rather
similarly developed in the two fossil forms but in P. ignavus the lamb-
doidal crest does not appear to be carried on by the squamosal, above
and lateral to the mastoid as in Pentapassalus.

The skull differences between Pentapassalus and Metacheiromys
(see Simpson, 1931) are similar to those between Pentapassalus and
Palaeanodon. However, the teeth in the Bridger genus are much
further reduced and the bullae are more completely ossified than in
Palaeanodon. The bullae in Pentapassalus resemble very much in
form those in Metacheiromys but are much more posterior in position.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN Si7/

I find no certain evidence in Pentapassalus of an entotympanic,
although a somewhat crushed zone around the posteromedial margin
of the left bulla is inconclusive, and the right bulla has been thrust
medially concealing this area. The glenoid surface for articulation of
the lower jaw is transversely much wider and has a better-developed
postglenoid process in Metacheiromys. The slope of the occiput is
similar in the two forms and the lambdoidal crests appear to be
similarly developed with respect to the mastoid exposure discussed
above. However, the braincase is much more inflated in Pentapassalus
and the skull is relatively shorter and broader posteriorly.

Comparisons with the Oligocene epoicotheriids show Pentapassalus
to be more closely allied to the Epoicotheriidae than to the Meta-
cheiromyidae, although known members of the latter family are more
nearly contemporaneous. P. pearcei strongly resembles the skull form
of the much smaller Epoicotherium unicum Douglass (1905, pl. 22,
figs. 13-16; see also Simpson, 1927, pl. 24). Most noticeable differ-
ences, however, lie in the dorsoventral attenuation of the rostrum, the
posterior elongation of the palate, the transversely much-constricted
choanae, and the continuation of the pterygoid crest onto the bulla
in Epoicotherium. The lambdoidal crest continues ventrally outside
of the mastoid portion, somewhat as in Pentapassalus, but the occiput
is much more inflated posteriorly in the Chadron form, strikingly as
in Chrysochloris, as has been noted by others. The dental formula
for the upper cheek-tooth series is the same in both fossil forms.

Xenocranium pileorivale Colbert (1942) from the still later Brule
beds of Wyoming is distinguished by the remarkable expansion of
the mastoid-squamosal region of the skull, much beyond that in either
Epoicotherium or Pentapassalus. The bullae in this form occupy a
posterior position but are transversely much elongated. Xenocranium
has lost one more postcanine tooth, above and below, than has Penta-
passalus.

Mandible.—The lower jaw of Pentapassalus pearcei is quite unlike
that in recent armadillos and Orycteropus. It corresponds closest in
size and form to that in Metacheiromys marshi Wortman (1903,
Pp. 347-352) or Metacheiromys tatusia Osborn (1904, p. 165; see
also Simpson, 1931, p. 303). The jaw is of nearly uniform depth
beneath the cheek teeth, deepening but slightly on the outside pos-
teriorly. Anterior to the single incisor the jaw tapers abruptly, and
a short distance posterior to the last molar the anterior margin of the
ascending ramus rises rather sharply to about go° from the line of the
tooth row, beneath its posterior deflection near the crest of the coro-

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

noid process. The coronoid process is much like that in Metacheiromys
and is separated by a broad U-shaped concavity from the condyle.
The condyle is gently convex, anteroposteriorly, and wider than long.
The angle is deflected markedly downward, below the inferior margin
of the horizontal ramus, and is widely separated from the condyle,
more so than in Metacheiromys. The masseteric fossa is a weak
triangular depression on the outer surface, and the inferior dental
foramen on the lingual side of the jaw has a markedly posterior posi-
tion in comparison with recent armadillos. Externally, a mental
foramen is exhibited beneath a point between the first and second
postcanine teeth and somewhat lower between the second and third.
The horizontal ramus reaches its thickest at the position of the canine,
and again at about the posterior margin of the last molar. The width
at the latter position is brought about by a shelflike ridge on the
lingual side, which in Pentapassalus can scarcely be attributed to a
pad as in monotremes, although this has been postulated for the more
nearly toothless Palaeanodon and Metacheiromys.

Comparison of the lower jaw of Pentapassalus pearcei with that of
Tubulodon taylori shows the latter to be less robust but with a little
larger teeth and the anterior margin of the ascending ramus rises
much more gently. The lower jaw of Epoicotherium unicum is not
known, but that for Xenocranium pileorivale is relatively much more
robust, though actually considerably smaller than P. pearcet.

Dentition—The dentition of Pentapassalus can be more clearly
1ecognized as the normal mammalian heterodont type than can mem-
bers of the Metacheiromyidae or other Epoicotheriidae. The presence
or absence of any upper incisors cannot be determined, but a single
lower incisor, separated from the canine by a small diastema, is
present. Strong upper and lower canines are preserved. These are
deep-rooted and the crowns are triangular in section, and enamel-
covered, except where worn through occlusion. The cheek-tooth
series includes five postcanine teeth in the maxilla and six in the lower
jaw. In upper series the first of these is close to the canine but
separated by a short diastema from the second. These appear to be
single-rooted premolars and are followed closely without diastema by
three molars. The first and second molars appear from their outline
at the alveolar margin to be three-rooted. The third upper molar is
more nearly circular at the alveolus but exhibits vestigial roots at the
base of the peg or stumplike crown.

There are three lower premolars, the first of which is very small,
and apparently pointed. It follows the lower canine by a diastema
which permits occlusion of the upper canine. The second and third

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 39

of the lower premolars are isolated by short diastemata, the last being
separated by a somewhat smaller space from the three uninterrupted
molars. The roots of the last premolar and second molar were in-
vestigated and found to be single ; whereas the first molar was found
to have two roots, which can be detected even well above the alveolar
margin. Presumably, M,; and the anterior premolars are single-
rooted.

The crowns of the cheek teeth are nearly flat with two planes of
occlusion meeting at a widely obtuse angle in a low transverse ridge,
generally near the middle of the tooth. The angle between these
occlusal planes is nowhere nearly so acute as in living armadillos.
In the premolar region these occlusal areas also slope more downward
and outward than in the molar region. A band of enamel surrounds
the crown or appears preserved on the lingual and labial margins, if
wear has not progressed too far. Occlusion of each check tooth is, of
course, with two in the opposite jaw, giving the crown a subdued
gable appearance. This is unlike the type of occlusion described for
Xenocranium.

The lower teeth in Pentapassalus are most like those in Tubulodon
taylori Jepson (1932), but do not exhibit the cuspate development
seen in the latter, probably a difference which may be attributed to
wear. The most significant difference lies in the persistence of double
roots in the last premolar and second molar as well as in the first
molar of Tubulodon. A superficial similarity is seen in the micro-
scopic tubular structure of the teeth of the two forms. This tooth
condition in Pentapassalus may not be the same as that in Tubulodon,
and is here apparently of no taxonomic importance, as it may be
observed in bones and teeth of other, quite unrelated forms in this
fauna.

Appendicular skeleton—The limb and foot material of Penta-
passalus pearcei is so completely like that in Metacheiromys that had
not the skull and jaws been found the material would have been
referred unhesitatingly to that genus. Of the various palaeanodonts,
limb material has been described only of Palaeanodon and Metachei-
romys. None so far has been recorded of the Oligocene forms. In
almost every instance differences noted from the more nearly con-
temporaneous Palaeanodon have been in the direction of M etachet-
romys. No special resemblance to Orycteropus limb material has been
noted. In all elements the limb material of Pentapassalus pearcet is the
same size or but a little larger than that in Metacheiromys tatusia
Osborn, as described and figured by Simpson (1931).

Scapula-—The right scapula is preserved, but incompletely along

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

the thin, prescapular border, and the acromion is missing. The ex-
hibited portion of the spine is remarkably high and rugged with the
crest turned posteriorly giving strength comparable to that of the
postscapular border.

Fore imb.—Comparison of the humerus (fig. 3a) is made with that
of Metacheiromys dasypus inasmuch as this bone in M. tatusia is not
illustrated ; nevertheless, correspondence in form is amazingly similar.
Differences noted in the illustration of P. pearcei and that of M.

Fic. 3.—Pentapassalus pearcei, new genus and species: a, Right humerus,
restored from left side, proximal, lateral, anterior, and distal views, natural
size; b, right ulna, anterior and distal views, approximately nine-eighths
natural size; c, distal portion of right radius, anterolateral and distal views,
approximately five-fourths natural size. Type specimen (U.S.N.M. No. 20028).
Upper Knight, lower Eocene, Wyoming.

dasypus, except for size, are due in part to orientation. The lesser
tuberosity extends somewhat lower on outer side of the proximal end,
and the width of the deltoid process distally appears slightly less, but
is incomplete in P. pearcei. The supinator ridge and much of the distal
extremity are almost identical in the two.

Only the distal portions of the radii (fig. 3c) are preserved. These
show an anterior crest which is well developed distally, but a short
distance (about 7 mm.) from the distal end it is abruptly reduced,
giving rise to a noticeable upward-directed process that is better de-
veloped in Metacheiromys. The distal articular surface shows no
distinctive features in comparison with the Metacheiromys radius.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 4I

The shaft of the ulna (fig. 3b) may be somewhat less curved
laterally in the anterior view than in Metacheiromys dasypus but the
olecranon is almost identical in its relative proportions and the inward
curvature of its extremity. The sigmoid notch is incomplete in
P. pearcei.

Pelvis and sacrum.—Portions of right and left innominate bones,
together with part of the sacrum are preserved. The right side is

Fic. 4.—Pentapassalus pearcei, new genus and species: a, Left femur, proxi-
mal, lateral, posterior, and distal views, natural size; b, left tibia, proximal,
posterior, and distal views, approximately five-fourths natural size. Type
specimen (U.S.N.M. No. 20028), Upper Knight, lower Eocene, Wyoming.

more nearly complete than the left, but both lack the pubic portion.
The ilium is slender and outwardly curved, extending, as in Metachei-
romys, well in advance of the sacrolumbar contact. Three elements
are involved in the sacrum as indicated by the fused spines, probably
about as in Metacheiromys; however, as shown in the referred speci-
men of M. marshi, there may have been a fourth centrum fused to
this series. The anterior zygapophyses for articulation with the last
lumbar are apparently much larger than in M. tatusia.

Hind limb—Both femora (fig. 4a) are included in the P. pearcei
skeleton, and these so closely resemble the femur of Metacheiromys

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. G7

tatusia figured by Simpson that no significant differences can be de-
tected. Comparison with Palaeanodon is fair, although Matthew’s
illustration of the femur in his figure 52 (1918), with his description
of P. ignavus, does not particularly resemble that shown in figure 65B.
In Dasypus novemcinctus the shaft is curved somewhat as in P.
pearcet but the greater trochanter is far better developed and the
relatively large third trochanter is more distally placed.

The details of the tibia (fig. 4b) are also decidedly like those of
M. tatusia. It is noted, however, in P. pearcei that the external
tuberosity of the proximal extremity extends a conspicuous process
laterally and distally carrying the articular surface for the fibula.
Also, the shaft may curve posteriorly somewhat more just beyond the
lower limit of the cnemial crest.

Only the proximal extremity of the right fibula is included in the
Knight material. This shows little of significance other than being
transversely flattened and the articulation for the tibia is a surface
almost at right angles to the longitudinal axis of the shaft.

Foot bones.—The foot material of Pentapassalus pearcei includes
the lunar, magnum, trapezoid, second and third metacarpals, and
various phalanges of the manus; but of the pes, only a part of the
astragalus is preserved. The lunar would seem to be much like that
described by Simpson for Metacheiromys dasypus ; however, it differs
from the illustration of this element in that the radial surface extends
entirely to the distal surface at the dorsal margin, much as in Dasypus
novemcinctus. The distal surface differs from that of the armadillo,
as described by Simpson, in that the articulation for the magnum is
sharply offset into two parts, the rectangular, more dorsal surface
being distinctly more distal and separated from the concave, nearly
circular, and more palmar surface by a sharp transverse ridge. A
slender, dorsoventrally concave articular surface is seen for the
unciform which widens somewhat palmad and distally. Dorsal to this,
and more laterally facing, is a facet for the cuneiform.

The trapezoid, in proximal view, is a triangular bone with its acute
apex directed palmad. The triangular surface for the scaphoid is
dorsoventrally concave in its palmar portion and convex dorsally.
A triangular surface is seen dorsally on the inner side for the tra-
pezium, and a plane crescentic surface dorsally on the external side
for the magnum. Distally, the trapezoid exhibits two dorsoventrally
elongate depressions, with an intervening saddle, the outer of which
is the longer, for articulation with the second metacarpal. The trap-
ezoid apparently does not articulate with the third metacarpal, nor
is this the case in Dasypus novemcinctus.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 43

The proximal surface of the magnum shows, as indicated above, a
rectangular shelf and raised, knoblike, more palmar portion for the
lunar articulation. The dorsal surface of the magnum is irregular,
much as in Metacheiromys dasypus, not so nearly rectangular as in
Dasypus novemcinctus. On the dorsal part of the inner aspect a plane,
pentagonal, somewhat proximal facing surface is seen for contact with
the trapezoid, and the palmar portion of the inner aspect exhibits a
rounded concave surface for a projection of the second metacarpal
much as in Metacheiromys. The outer surface of the magnum exhibits
an irregular concave surface for the unciform. The distal articular
surface is a dorsoventrally, deeply concave saddle for the third meta-
carpal.

The second metacarpal is a very short and sturdy bone, about three-
fourths as long as the third metacarpal. The proximal extremity is
nearly rectangular, deeply grooved dorsoventrally in the more dorsal
portion, and the inner-palmar angle is formed by a prominent rounded
knob separated from the articular surface. The dorsal portion of the
inner margin shows a small, inward-facing surface for articulation,
presumably with the trapezium, and on the dorsal portion of outer
margin the facet for the third metacarpal faces somewhat distally.
The outer more palmar surface for the magnum is relatively smaller
and is not oriented so distinctly proximal as in Dasypus novemeinctus.
Distally, the surface for articulation with the first phalanx is much
broader than the proximal extremity, and is dorsoventrally more con-
stricted on its inner side than on the outer. Dorsally the surface is
transversely cylindrical with prominent, symmetrically placed pockets,
palmar to the convexity. The shaft of the second metacarpal is very
broad dorsally, this aspect being much emphasized by the process for
the extensor muscle, which is about midway on the outer margin.

The proximal extremity of the third metacarpal is much more
convex, dorsoventrally, than the second. The dorsal portion is rela-
tively broad and concave for articulation of the magnum. A short dis-
tance palmar to the dorsal portion the proximal extremity is notice-
ably constricted transversely, giving the proximal aspect a T-shaped
appearance. The external knob of the dorsal expansion articulates
with the second metacarpal, whereas the internal knob shows a nar-
row, limited surface for articulation with the unciform, very much
less than in D. novemcinctus, but immediately distal to this more
proximal surface is a relatively large, outward and palmad-facing con-
cavity, faceted for articulation with the fourth metacarpal. The distal
articular surface is for the most part transversely cylindrical with the
palmar pockets more feebly developed than in the second metacarpal.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS LL,

The shaft is sturdy and slightly arched dorsally. The process for the
pars brevis of the extensor muscle is very prominent, proximal to the
midpoint of the inner margin of the dorsal surface, but somewhat
distal to the equivalent process on the second metacarpal. The latter
condition is reverse to that seen in Metacheiromys dasypus.

The metacarpals and phalanges are not only smaller but relatively
shorter than in Palaeanodon ignavus. They resemble materials from
the Lost Cabin beds attributed by Matthew to ?Palaeanodon, species,
but are rather smaller. These Lost Cabin elements might, of course,
represent Pentapassalus or Tubulodon.

The intermediate phalanges and claw cores are quite like those
figured for Metacheiromys, except that there is a suggestion that the
articulation on the claw may extend somewhat more dorsoventrally
around the distal articular surface of the second phalanx than in the
Bridger type and the claws may be more curved in a vertical plane.

The head of the astragalus, which is the only part of the hind foot
preserved, shows a surface broadly curved transversely for the
navicular, and a relatively large, slightly convex sustentacular facet
close to that for the navicular. The details are much like those shown
by Matthew for Palaeanodon ignavus. The body of the astragalus is
missing.

Remarks.—Undoubtedly the most significant, and certainly the
most interesting find of the several seasons’ collecting in the Knight
beds is this rare edentate mammal. During the earlier stages of this
investigation a rather striking tubular structure noted in the teeth was
thought to ally it peculiarly with Tubulodon from the Wind River
beds, and to place it in some position of relationship to Orycteropus.
Further observations, however, have shown that so far as the Knight
materials are concerned that is not the case. The microscopic, irregu-
lar, anastomosing clusters of tubules, quite independent of the Haver-
sian system, were also noted in otherwise dense and somewhat trans-
lucent bone and teeth of certain other forms in the fauna, cautioning
against regarding the structure as significant in teeth of Pentapassalus.
The structure noted in the above material was surely of post-mortem
development.

Pentapassalus shows very little of importance in the skull structure
that would tend to ally it to the Tubulidentata, no more than edentates
in general might resemble them. In its general skull form and in
retention of the teeth it is most closely allied to the Oligocene Epoico-
therium of the various palaeanodonts, and is placed in Epoicotheriidae
rather than with the longer-skulled, more nearly edentulous and more
nearly contemporaneous metacheiromyids. Limb and foot structures

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 45

are similar to those in Metacheiromys but it is very likely that Epoico-
therium was similarly adapted.

MEASUREMENTS OF TYPE SKELETON OF

Pentapassalus pearcei, NEW GENUS AND NEW SPECIES, U.S.N.M. NO. 20028
Skull: aan
Length, anterior extremity of nasals to condyles.................... a59.0
Wace at postoroitel COnstriction'.-.).':....'s.....s. eee dee deen tee a 18.0
Length of cranium from postorbital constriction to lambdoidal crest.. 23.5
Width of cranium across squamosal portions of lambdoidal crests.... 34.7
Depth of occiput from top of crest to bottom of condyles........... 19.2
Mandible:
Length of mandible from anterior extremity to condyle............. 42.5
Wepthrar jaw beneath Ms internally. 3.2 o.c.'s cadets cee cad oe ew eee 5.2
Thickness of jaw at widest part immediately behind M;............. 4.0
Dentition :
aM ee Cle ees ce ea ee aoe Ale Sele oh hen Hoag otis uo eee 15.7
HL UO) Wiles « OS Ae SiG caches SRNL pCa RRC oe CREPE Ta nn inte Eman URIS PAE
ROCCO a i ad Neves et Con hy Pet ieee ansi's. stn '& Sivisnerone <n los do iaies ic Fane enna 18.5
Tae VES Roma ten, wom meee) A wll ara an Fe ls tera ioe (43 Ried ome eae 8.5
NEC Marae a Nurse. eta, thei s titcidid wahiald’s vale Med. ok sla @e «eee 6.9
Anteroposterior diameter of last upper premolar................... 2.0
Amreropostenion diameter .OL, NAA. ics seisuis saa es o's d ecw on coe ele snes 3.0
Aer HEVCRSORIAIeLEl li N=. se tera 2a shige a ciahass ciedersre wis o/s. ofa ceases 2.5
Anteroposterior diameter of last lower premolar................... ae
sbransyerse; diameter o£ last; lower premolar....0.....4--a0sssseene 1.6
AMLerONOstemMOlma@iatneteniOn Niocas +. as sce sisielc «3 ate oto eked yn serene 2.4
siransverse Ciaineter Of Win. ct tert cc ses e ce bese sls cateses cede stage mere 1.9
Fore limb:
Length of humerus, head to articular surface for ulna.............. 37.0
Transverse width of humerus across proximal tuberosities.......... 12.1
Greatest transverse width of humerus across condyles of distal
SEMEN Agama tay ee os SA aro Sieh a or Bea ahie os doa) a oie ac aiafoie Sre, aves eyo S EST sp cle cme are tne 21.7
Depth of distal extremity of humerus across capitulum for ulna...... 5.5
Length of deltoid crest from greater tuberosity................-.00. 28.0
Length of supinator ridge from surface for ulna................... 22.0
Wet Gi distal Cxiceinity, Of TACUS 02:0. <6: 66 = eos aot eels ea ameins 7.0
Depth of distal extremity of radius incl. longitudinal ridge.......... 75
RETEALESEMEDSEN OG! MINA! Loe esc ee ose ches oe Fee ces Bam olde oe eelnneNys 48.7
iength ef.olecranon: from- sigmoid: notch: ..is.00.5 duce ae sean ne: a 18.5
Greatest) width of distall extremity of ulna. /....-2---0-++.c0---- 6.0
ecto ObeSeeand) etACAL DAL o/s sions lax di aikis a/os-e\ oe ais @ epee a te Sie te wT
Width of proximal extremity of second metacarpal................. 3.5
Width of distal extremity of second metacarpal.................... 5.0
MPEM EMO CHING METACATDAl 4. 3 4-5 <ja.a0 aieje cpa eye, sacar maiuishnerey eine sieheye Wale 0.5
Width of proximal extremity of third metacarpal.................. 5.5
Width of distal extremity of third metacarpal..........s0eseeeeeees Bea

a, Approximate.

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Ti7

Hind limb: mm.
Greatest’ lenothxok, femur i seeds vars Sse tapa cine one aie cle sis eke actos eee 50.0
Width of proximal extremity of femur across head and greater

{5 MOL COTO dc) UMAR ESCA ee SI RINE OL Rn EET Gee cts o MOIR eOIy chats aeL0 6 15.0
Width across condyles of distal extremity of femur.................. 13.5
Depthtor distalvextremityFonaemute eres a scree eter 10.5
Width ‘of proximal extremity jon dibia’s <5 <cctem ereciescueia + ate ete al reat 12.4
Depthvot proximal extremity or tibideanre scree ate eee aoe 10.0
Width of distal vextremity of tibia..62.c6 ae eitsoe on sete eae 9.4
Deptheonadistalwextiem tye otatibiaerermeemucmeeeeecic eerie 5.6
Woadthvoteproxtmalsextremibvionitibtlanemanecern ener r icine ncr ttre a2
Depthict proximal wextremitylor tibiulanen uae aseee erratic cites 6.6

RODENTIA
ISCHYROMYIDAE

PARAMYS, cf. COPEI 1? Loomis

Seven isolated lower-jaw portions and an isolated upper molar,
in addition to a specimen, U.S.N.M. No. 19306, including both rami
of the mandible with some associated limb and vertebral fragments,
represent in the La Barge fauna a moderate-sized paramyine rodent,
tentatively referred to the species Paramys copei Loomis (1907a,
p..128).

Examination of the teeth of the upper Knight paramyine jaws in
comparison with various specimens of Bridger Paramys shows little
or no significant structural differences upon which, on teeth alone, one
could reasonably separate these upper Wasatchian forms generically
from true Paramys. The lack of definitive characters in the teeth was
noted by Matthew (1920) in his description of Reithroparamys.
Paramys appears to be a highly diversified genus, with Wasatchian
species assigned to it more than covering the possible size ranges.
Moreover, minor differences in teeth attributed to some of these species
are probably, in view of Bridger variations, of no systematic signifi-
cance, so that several of the species are likely invalid. Final allocation
of the Knight species herein discussed awaits revision of the para-
myids currently under study by A. E. Wood.

18 While this paper was in press, Dr. A. E. Wood, who is revising the
paramyids, examined these specimens and informs me that most of them surely
represent Paramys copei, but that U.S.N.M. Nos. 19305 and 19306 should be
referred to Paramys major, and a third, No. 19307, may be a new species.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 47

PARAMYS, cf. BUCCATUS !4 (Cope)

An immature lower jaw, U.S.N.M. No. 19302, with the molar teeth
broken off, but with the last premolar erupting, is compared, on the
basis of the size of Ps, to Cope’s (1877, p. 171) species, Paramys
buccatus. There are also in the upper Knight collection fragments
of a skull, including a single upper molar which cannot be distinguished
from the type of this New Mexican species. Although Loomis (1go7a,
p- 130) considered this species as possibly belonging to Sciuravus,
as Matthew (1910, p. 51) has noted, it is clearly paramyine and not
sciuravine.

A left ramus of a mandible with portions of Ps to Mz, inclusive,
No. 12845 of the Knight collection of the American Museum, is very
close in size of P, to U.S.N.M. No. 19302, and may also represent
Paramys buccatus. This was included among the materials collected
by Granger and Miller in 1906 from the bluffs along Bear River near
Knight station.

SCIURAVUS, possibly S. DEPRESSUS Loomis

Recognition of Sciuravus in the lower Eocene has in the past been
attended by serious doubt. In 1907 (a, p. 130) Loomis described three
upper molars and a lower molar as the species Sciuravus depressus,
from a locality in the Wind River beds near Lost Cabin, Wyo.
Matthew, though cognizant of the occurrence in 1910, as shown by
his phylogenetic chart and distribution table, did not comment on it,
and in 1918 (p. 619) ignored the possibility, and listed Sciuravus
without comment only in the middle and upper Eocene. Troxell in
1923 (a, p. 385) dismissed the possibility with the comment that from
the drawings he judged the form to be not Sciuravus but a separate
genus. Wilson (1938, p. 129) in his revision of Sciuravus based on
the Bridger rodents in the Marsh collection cited the occurrence but
added Troxell’s comment. Again in 1949 (p. 79) Wilson cited
“Sciuravus depressus’ in the Lysite fauna, with the reservation that
although this species apparently could not be referred to the genus
Sciuravus it might be tentatively included in the Sciuravinae. No
critical examination of the specimen has been reported since Loomis
described it, and Loomis’s illustration is clearly unsatisfactory, so
that in reality the presence of Sciwravus in the lower Eocene has not
been entirely discredited.

14Dr. A. E. Wood has since examined these specimens and indicates that
this small Wyoming Paramys should be referred to Paramys excavatus Loomis
rather than the New Mexican P. buccatus.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IT7

Sciuravus material in the upper Knight collection consists of the
left ramus of the mandible, U.S.N.M. No. 19309, with P, and M,,
from about 12 miles north of Big Piney, Wyo. The species repre-
sented in the La Barge fauna is intermediate in size between Sciuravus
nitidus Marsh and Sciuravus bridgerit Wilson of the Bridger fauna.
It appears to be close in size to Sciuravus depressus, judging by
Loomis’s figure of 8 mm. for the lower molariform series.

TILLOMYS SENIOR,!* new species
Text figure 5

Type.—Right ramus of mandible with P, to Mg, inclusive, U.S.
N.M. No. 19308.

Horizon and locality—Upper Knight beds, La Barge fauna, SW}
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

No. 19308), type specimen; occlusal view, 6; lateral view, 3. Upper
Knight, lower Eocene, Wyoming.

Specific characters ——Size much smaller than Tillomys senex Marsh
(1872, p. 219) and appreciably smaller than Tillomys parvidens
(Marsh) (1872, p. 220). Teeth with cusps very low and crests
subdued, increasing in size from P, to M,. Tooth row shorter than
in T. parvidens, but jaw relatively deeper. Masseteric fossa does not
extend so far forward with respect to teeth as in T. parvidens.

Description—The above type is the only known specimen but is in
a rather good state of preservation. The crowns of the first three
cheek teeth are intact though worn to the point of exposing dentine
in the central portion of each cusp. The occlusal portion of the incisor
and the crown of M; are broken off.

15 Older, comparative of senex; forerunner of Tillomys senex.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN AQ

The details of the cusp arrangement in Tillomys senior are so like
those in T. parvidens that confusion with other known genera of
Eocene rodents seems entirely improbable. The cusps are even more
subdued, possibly in part due to wear, than in the Bridger material
examined, particularly the metaconid, and the cusps are perhaps more
conical with less tendency toward development of crests. The ento-
conid is isolated with a very weak, posteroexternally directed crest,
distinct from the internally directed posterolophid. The mesoconid
is a simple conical cusp wedged between the protoconid and hypo-
conid. The pattern of wear on the protoconid suggests two low spurs
directed toward the metaconid. Each of the three preserved teeth
shows the trigonid to be narrower than the talonid and each succeed-
ing tooth is wider than that preceding. Ms, though broken off, is
clearly longer than M., but the width cannot be ascertained.

The teeth in Tillomys senior are only slightly larger than in the
type of Mysops kalicola from the Lost Cabin beds, but the pattern
of the crown is essentially different. M. kalicola in addition to having
highly rugose enamel in the more deeply basined crown has sharper,
more arcuate cusps and the crests or lophs are developed more con-
tinuously across the crown. Moreover, the jaw of M. kalicola, while
deeper beneath M;, becomes shallower forward beneath the anterior
cheek teeth, and is transversely more slender.

MEASUREMENTS OF TYPE LOWER JAW OF
Tillomys senior, NEW SPECIES, U.S.N.M. NO. 19308

mm.
engthtonscheek-toothwsenies. ese ca-ses aa nn eeieeietn omer tr a 6.6
Tieereeet Uaericath clotat sti Mia aad so cg leh ci defen cyin's Bash rare sdgayoumtey a pestmnamtrerae 4.6
Ps, anteroposterior diameter: greatest transverse width...... Tsien.
Mi, iB of ai ais eran os P55
M2, % 5 +e “ aay eee TOL EF
Depth of jaw beneath Mi, on inner side..............cccecees 5.9

a, Approximate.

Remarks—tThe occurrence of Tillomys senior in the Knight fauna
of the Green River basin extends the known geologic distribution of
Tillomys into the lower Eocene. The principal significance of this
fact is the additional evidence afforded of the remarkable conservatism
of rodent genera in Eocene time, adding to that of Paramys and
Sciuravus. This is in striking contrast to the still more remarkable
evolutionary change which took place in certain other groups of
mammals between Lost Cabin and Bridger time. All of which is
further proof of the antiquity of the order Rodentia, and arouses
speculation as to why so little rodent material is known from the
Paleocene, and where their earlier center of dispersal might have been.

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

CARNIVORA
ARCTOCYONIDAE
THRYPTACODON, near T. ANTIQUUS Matthew

Although Matthew (1915a, p. 7) stated that Thryptacodon has not
been found in the Lysite or Lost Cabin, Van Houten (1945, p. 452)
noted the presence in American Museum collections of a specimen of
T. antiquus labeled “?Lysite” from the Big Horn basin. The upper
Knight materials, which consist of a right upper molar and a tenta-
tively included maxillary fragment with two milk teeth, appear to
extend the range to Lost Cabin time.

The isolated upper molar, U.S.N.M. No. 19476, corresponds closely
to Thryptacodon antiquus material of Gray Bull age in the National
Museum collections and is almost identical in size and pattern to the
second molar in the type specimen figured by Matthew (1915a, fig. 4).
The maxillary portion, U.S.N.M. No. 19343, with the two teeth
believed to be deciduous third and fourth premolars, rather resemble
P* and M?* of Thryptacodon, but are noticeably elongate anteropos-
teriorly, particularly the labial portion. The appearance and propor-
tions of these teeth are about as one might anticipate in a Thrypta-
codon milk dentition.

MESONYCHIDAE
PACHYAENA? species

Two specimens, a single lower tooth and one associated with bone
fragments, would appear to represent Pachyaena in the New Fork
fauna. The development of the paraconid on these suggests P, and
M,. They are rather close in size to these teeth in Pachyaena gracilis.

OXYAENIDAE
OXYAENA, species

A single lower molar, obtained by Princeton University in associa-
tion with the Ambloctonus jaw described below, and carrying the
number P.U. 16176, was among the Knight materials which Dr.
Jepsen has permitted me to include with my study of the Smithsonian
collections. This tooth is readily recognized as a first lower molar
of Oxyaena; however, the species represented is not evident. In size
it corresponds very closely to the New Mexican Oxyaena lupina Cope.
Resemblance on the basis of such limited material, however, scarcely
justifies extending the range of this species to the late lower Eocene
of Wyoming.

no. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 51

AMBLOCTONUS, cf. MAJOR Denison
Plate 6, figure 2

An Ambloctonus lower jaw, obtained by Princeton University
from a locality in the upper Knight about 2 miles southeast of La
Barge, Wyo., was kindly loaned to me for study by Dr. G. L. Jepsen.
This specimen, P.U. No. 14720, includes P. to P4, inclusive, and Ma,
and exhibits alveoli for the canine and M,.

The premolars of the Princeton specimen are a little like those in
Oxyaena forcipata, but, as in Ambloctonus, are distinctly broader,
particularly across the talonid, perhaps blunter, and P, exhibits a
better-developed parastylid. These teeth are relatively longer and
apparently less hypsodont than in Palaeonictis. M, was remarkably
large, as indicated by the alveoli, the posterior of which is consider-
ably longer, anteroposteriorly, than the anterior. Mz, is of reduced
size, with a much curved shearing blade, no metaconid, and a very
small talonid. The narrowness of the talonid and absence of a meta-
conid on this tooth are characteristic of Ambloctonus, and distinguish
this form from Palaeonicits.

The type of Ambloctonus major Denison (1938, p. 176), as figured
by Matthew (1g15a, fig. 53) under reference to Ambloctonus hyae-
noides, consists of a fragmentary jaw with M, and part of P,. Direct
comparison in not feasible, but the size of the preserved molar in the
Lost Cabin type specimen corresponds almost exactly to the space
for the missing tooth in the upper Knight jaw.

Among fragments of carnivore teeth found in the New Fork beds
is a left P; which resembles closely this tooth in the Princeton jaw
from near La Barge. It is about the same length but slightly wider.
It lacks a paraconid as in both Ambloctonus and Palaeonictis but its
profile in lateral view is much more like Ambloctonus.

LIMNOCYONIDAE
PROLIMNOCYON ELISABETHAE,!® new species
Plate 5, figures I-3

Type.—Both rami of the mandible with the canine and P, to M2,
inclusive, represented, U.S.N.M. No. 19350.

Horizon and locality—Upper Knight beds, La Barge fauna, north
side of “Red Hill,” western margin of SW4 sec. 23, T. 29 N., R. 111
W., about 5 miles southeast of Big Piney, Sublette County, Wyo.

16 Named for my wife, who found the type specimen.

iS)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

onl

Description —Teeth a little smaller and much more slender than in
Prolimnocyon antiquus. Pz to Ps with increasingly prominent para-
style. Talonid of molars narrower than in P. antiquus and lower jaw
a little shorter, shallower, and much more slender. M3 single-rooted.

Discussion—In addition to the pair of jaws described above as the
type of this species, there are portions of right and left maxillae of one
individual, U.S.N.M. No. 19348, including P* to M? between them,
and a third individual, U.S.N.M. No. 19349, represented by a left
mandibular ramus with portions of P, and M,.

The type of Prolimnocyon antiquus from the Lost Cabin beds con-
sists of right jaw portion with only the heel of M, preserved. An
isolated M, from the opposite side is included with the type and may
have been used by Matthew (1915, fig. 63) to illustrate this tooth
in the jaw. The M, referred to the type is fairly robust and has a
decidedly broad, basined talonid, wider than in M, and relatively
much larger than in the type of the earlier Prolimmnocyon atavus. M,
in the type of Prolimnocyon elisabethae is smaller, much more slender
and has a considerably smaller talonid than that attributed to P. anti-
quus. The premolars in P. elisabethae are almost as long as in P. antt-
quus but more slender as indicated by root portions in the type of the
latter. A referred specimen of P. antiquus, A.M. No. 2971, shows
P; and P, to have almost no parastyle, and the parastyle is weak or
absent in material of Gray Bull P. atavus. In P. elisabethae the para-
style is weak on P, but decidedly prominent on P,.

Right and left maxillae, U.S.N.M. No. 19348, have between them
P* to M?, inclusive. A single small alveolus is preserved of M® but
the specimen is incomplete lingual to this point so that the presence
of additional alveoli for this much-reduced tooth cannot be deter-
mined. P* shows a single large primary cusp and a sharp high-cusped
deuterocone, the latter of which is separated by a deeply indented
anterior border from the distinct parastyle. The tritocone is somewhat
elongate and approaches the crested form characterizing the first
molar. In M* the paracone and matacone are distinctly more separated
than in Prolimnocyon atavus of the Gray Bull stage, and the isthmus
to the protocone is a little more constricted, with a better-developed
and nearly continuous cingulum around the protocone. Also, the
parastyle consists of a complex of two small stylar cusps together
with a third small cusp between these and the paracone. M? is, of
course, transverse with a greatly elongate parastyle, no metastyle, and
a vestigial metacone. The striking difference between this tooth in
Prolimnocyon and that in Sinopa is one of function, and Butler (1941)
has shown in a study of Hemiucentetes something of the mechanism

No. 18 LOWER EOCENE KNIGHT FORMATION—-GAZIN 53

or explanation of the transition that may account for the ways and
means of divergence between these two creodonts. However, such
a separation obviously took place at a much earlier and unspecialized
stage of dental development in both lines, and by lower Eocene time
this divergence has a basic, nonadaptive significance.

The numerous characters that have been used (Denison, 1938,
Ppp. 241-242) to support transfer of the Limnocyoninae to the Hyaeno-
dontidae are for the most part matters of degree in distinction from
the Oxyaeninae, and largely of adaptive significance in comparison
with Proviverrinae. Demonstration of a probable common ancestry
does not necessarily justify inclusion of two groups of genera in the
same family, particularly when these related groups diverge and
develop their own taxonomic complex, such as horses and tapirs.
Certainly the genera most typical of the family Hyaenodontidae, those
included in the Hyaenodontinae, are strikingly distinctive and unlike
Limnocyon and its allies. All things considered I find inclusion of the
Limnocyoninae and Machairoidinae in the Hyaenodontidae untenable.
On the other hand, the probable origin of Prolimnocyon, or rather
of its ancestry, would suggest, although this is uncertain, a poly-
phyletic Oxyaenidae were the Limmnocyoninae to be retained in that
family.

HYAENODONTIDAE
SINOPA VULPECULA Matthew

U.S.N.M. No. 19347, which consists of maxillae and jaws together
with fragments of other skeletal portions, is regarded as representing
in the La Barge fauna the small Lost Cabin species, Sinopa vulpecula
Matthew (1915a, pp. 80-82). The teeth are close in size to those in
the type but the jaw is slightly shorter, shallower, and straighter, also
the coronoid portion not so high. P, is absent in both rami, but a
slightly rugose depression well ahead of P, suggests an earlier pres-
ence, but loss before death. Though the animal was evidently not
an aged adult, the third molar in both rami was broken to root stubs
which were well worn before death. Also, the first molar is missing
on the right side and the anterior alveolus for this tooth was com-
pletely closed before death.

The upper teeth are undamaged, so far as represented, and in the
right maxilla P? to M® are present. Wear in the molar series was
evidently restricted for the most part to occlusion with M, and the
posterior root of Mg, although earlier occlusion with the trigonid of
M; is indicated. However, wear resulting from M, is very slight,

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

indicating a very early loss of this tooth. The upper molars of No.
19347 are much larger than those in the Lysite specimen figured by
Matthew as possibly representing Sinopa vulpecula, They are close
in size to those of the type of Sinopa multicuspis from the Eocene of
New Mexico. The most noticeable difference from the latter is a
greater lingual extension of the protocone portion in the molars and
possibly less development of the parastyle on M? of the upper Knight
specimen. The slenderness and lingual projection of the protoconal
portion is rather noticeable in comparison with middle Eocene forms,
such as Sinopa rapax.

It is also noted that the upper Knight form may compare favorably
in size with Tritemnodon? whitiae Cope, but the lower premolars are
more robust and longer in the latter, and the talonids of the lower
molars are smaller. Upper teeth of a referred specimen of T.? whiitiae,
A.M. No. 4781, show no metacone on M®* which is present in the
Knight material, and P* shows a small talon not seen in the Knight
specimen referred to Sinopa vulpecula.

SINOPA, cf. STRENUA (Cope)

A single left maxillary portion with the last two molars, U.S.N.M.
No. 19345, tentatively referred to Sinopa sirenua, shows the presence
of a larger species of Simopa in the La Barge fauna than is repre-
sented by the better-preserved material described above as belonging
to Sinopa vulpecula. Direct comparison with the type of S. strenua
is not feasible, inasmuch as the New Mexican specimen includes only
the lower jaws; however, Matthew (1915a, pp. 74-75) has suggested
the possibility of this species being represented by various Lost Cabin
specimens. On the basis of size alone No. 19345 might come within
the range of S. strenua.

MIACIDAE
DIDYMICTIS ALTIDENS Cope

Matthew’s (1915a, pp. 19-26) revision of the lower Eocene species
of Didymictis is an oversimplification of the situation, and includes
errors which have led to confusion in attempts to identify Wasatchian
materials representing this genus. It seems unlikely that he examined
the type of Didymictis protenus, as the actual measurements of it are
less than the range which he has given for the subspecies of this name.
For the record these are P,-Mz, inclusive, 53 mm.; and M,-Ma,,
18.3 mm. The ranges given for Didymictis protenus lysitensis and
Didymictis altidens would appear to be diagnostic for the Lysite and

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 55

Lost Cabin horizons, whereas this is not the case. Several specimens
of Lost Cabin Didymictis in the National Museum collections are
within the range given for D. p. lysitensis. This is also true of
the upper Knight materials, of which there are some 14 specimens
from the vicinity of La Barge and Big Piney, Wyo. I do not believe,
however, that a Lysite age is indicated thereby. It seems more
probable, as Simpson (1937, p. 15) has suggested, that Didymuctis
protenus lysitensis is either a synonym of D. altidens, or a subspecies
of the latter rather than of D. protenus. On this basis I have referred
the upper Knight materials to D. aliidens. They are scarcely com-
parable to the type D. protenus and only a little smaller than dimen-
sions given by Cope (1880a, p. 746) for D. altidens.

Much of the Didymiciis material in National Museum collections
from the Gray Bull exhibits somewhat smaller proportions than
D. protenus, and should any prove to represent a distinct species,
the name Didymictis leptomylus, as well as the larger D. curtidens, is
available. Simpson has shown that D. leptomylus possesses a remark-
ably small M,; however, he regards the locality for the type as in
doubt. This was clarified by Wortman, as reported by Matthew
(1915b, p. 314), subsequent to Matthew’s work on the Wasatchian
carnivores. Hence, Didymictis leptomylus Cope, Haplomylus, speiri-
anus (Cope), and Esthonyx spatularius Cope, originally regarded as
from Wind River beds, are shown to be Gray Bull, and almost surely
lower Gray Bull. This is in accord with locality corrections later made
by Cope without explanation.

VIVERRAVUS LUTOSUS,!7 new species
Text figure 6

Type.—Right ramus of mandible with Pz to M4, inclusive, U.S.
N.M. No. 19339.

Horizon and locality —Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description—Size of teeth intermediate between Lost Cabin
Viverravus dawkinsianus (Cope) and lower Gray Bull Viverravus
acutus Matthew, but depth of jaw shallow as in V. acutus. P, two-
rooted, P; with posterior accessory cusp, but less well developed than
in P,. Talonid of M, with trenchant, external hypoconid, and lingual
portion distinctly basined.

17 From Latin lutosus, muddy, with reference to Muddy Creek where the
type was found.

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Remarks.—Viverravus lutosus material includes approximately
eight specimens, all lower-jaw portions except for an isolated P*.
Most of these come from a badland locality on Muddy Creek about
12 miles north of Big Piney. The species is probably closest to
Viverravus dawkinsianus, particularly in the development of the
anterior premolars. However, it has distinctly smaller teeth, par-
ticularly Py, and the lower jaw, though fully mature, is as shallow as
in Viverravus acutus. The shallowness is most noticeable posteriorly
beneath the molars, with the greatest depth beneath P,. In other
Wasatchian species the depth apparently increases posteriorly beneath
the molars.

Hoe

Fic. 6.—V iverravus lutosus, new species: Right ramus of mandible (U.S.N.M.
No. 19339), type specimen; occlusal and lateral view, X 2. Upper Knight, lower
Eocene, Wyoming.

MEASUREMENTS OF TYPE LOWER JAW OF
Viverravus lutosus, NEW SPECIES, U.S.N.M. NO. 19339

mm
Length of cheek-tooth series at alveoli, Pi-M2, incl............ 237;
Length of premolar series; Pi-Pa 1Cl..-.. ce seedee es seo cen. 15.4
ensthvorimolar series, Mi-Marinch a.guces...s+-ses52 oceans 8.7
Tength of sPe ci widthwotePatess denueeec atone eee AiO)R Milas!
engthivior My sswidthy or Misc wich ss sawed cents meee eee Bel 20
Depth ior yaw beneath Pisiseecs oon ese toe eee oe 5.4
Depthviorsyaw beneath? Note wiih wee ties crecenleee icine amen 5.3

UINTACYON ASODES,'® new species
Plate 6, figure 1

Type.—Left ramus of the mandible with the canine, P4,, and M,
preserved, U.S.N.M. No. 19351.

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

18 From Greek ao#dns, muddy, etc., with reference to Muddy Creek where
the type was found.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 57

Description.—Size near that of Uintacyon massetericus (Cope),
but jaw a little deeper, and teeth wider and more robust. Premolars
uninterrupted, except for very small diastemata before and after Py.
P, without paraconid. M, about 4o percent longer as measured at
roots. Mz; single-rooted.

Remarks.—Uintacyon asodes is represented by a single specimen
which consists of a lower jaw in which are preserved only the canine,
last premolar, and first molar. The anterior premolars and posterior
molars are indicated by root portions or alveoli. P, and M; are single-
rooted. Comparison of Uintacyon asodes with the referred specimen
of Uintacyon massetericus, which Matthew figured (1915a, fig. 24)
instead of the type, shows the carnassial to be wider and the premolars
to be less reduced and in closer sequence. Also, the lower jaw is
shorter and deeper and would appear to have a straighter inferior
margin or profile ; however, the anterior part of the specimen Matthew
figured is damaged and the restoration may not be correct.

Uintacyon is rare in lower Eocene deposits and this is apparently
the first record of its occurrence in beds of Lost Cabin age. Repre-
sentation in later Wasatchian was to be expected, however, inasmuch
as the typical and better-known species are Bridger middle Eocene in
age. Uintacyon asodes shows a definite trend toward the latter forms.

MEASUREMENTS OF TYPE LOWER JAW OF
Uintacyon asodes, NEW SPECIES, U.S.N.M. NO. 19351

mm,
Length of dentition from P: to Ms, incl., at alveoli.......... a 35.0
Length of premolar series, P: (alveolus) to Ps.............. 18.1
Length of molar series, Mi to M; (alveolus)................ a 17.0
Benotvote bar hwidthinole lat sc wei cir sien +. cfeleeis.< a1 s/n ee eloiatale caecs OEtins ys
Length of Mi: width ‘of M: at trigonid....5.... 000. cece eles FAB Be
Hens thwotelswaburOOUSm nays wrstite sees oF toa ieseieieinie ace aferw Wiovepelorere 5.9
Depth of jaw beneath Ps, lingual side.............cc.eeeeees 13.0
Depth of jaw beneath talonid of Mu, lingual side............. 12.2

a, Approximate.

MIACIS, cf. LATIDENS Matthew

Lower jaw portions of three individuals, all from 12 miles north
of Big Piney, exhibit teeth which are more like those in Miacts than
any of the other miacids. One of these specimens, U.S.N.M. No.
19335, includes portions of both rami and has preserved all the cheek
teeth, except P,, on the left side. The measurements of these are
almost the same as those given by Matthew (1915a, p. 34) for the
type of Miacis latidens. Unfortunately, however, the type exhibits
only the last two molars, so that only limited comparisons can be

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES E17

made. The upper Knight jaw has about the same depth beneath P,
or P., but is much deeper beneath the molars, particularly Ms. M,
corresponds in detail to that in the type and is about the same size;
however, M; would appear to be slightly more like M, in outline than
this tooth does in the type.

In addition to the few comparisons permitted with the type, it is
further noted in the upper Knight specimen that P, and P, are isolated
by diastemata, but that P; to M; are in a closed sequence. Of the
premolars, only P, exhibits an accessory posterior cuspule. The
trigonid of M, is high but those of the following molars are much
reduced. The talonid in each of the molars is basined, though it ap-
pears somewhat more trenchant, particularly M2, than in Vulpavus
australis, and each tapers somewhat more posteriorly than in the latter.
Moreover, M; appears more reduced than in Vulpavus, and although
the crown seems to be a reduced image of M,, the root portions are
not so well separated. In U.S.N.M. No. 19334 the alveolus appears
to be a single bilobed opening. The distinctions between species of
Miacis and Vulpavus, though better defined in the middle Eocene, are
less clearly discernible in lower Eocene materials, and some of the
more fragmentary specimens, including isolated teeth, in the Knight
collection could not be assigned with any certainty.

VULPAVUS ASIUS,!9 new species
Plate 5, figure 4

Type.—Portions of both rami of the mandible with P; and M,-Mg,
U.S.N.M. No. 19337.

Horizon and locality——Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Description——Depth of jaw about as in contemporary Vulpavus
canavus Cope but teeth closer in size to those of /ulpavus australis
Matthew of the New Mexican San Jose beds. The first two molars
are of about the length of those in V. australis but relatively wider,
and P, and M,, as indicated by root portions, are longer. The trigonid
of M, is higher, although relatively not so high as in Miacis, and the
paraconid and metaconid are less widely separated giving the “V”
of the trigonid a more acute angle at the protoconid.

Discussion.—V ulpavus asius may be represented by some half
dozen specimens encountered at various localities in the upper Knight

19 From Greek dovos, slimy, muddy; with reference to Muddy Creek where
the type specimen was found.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 59

beds. These appear to represent a form having a jaw about as robust
as Vulpavus canavus, but with teeth significantly smaller. In this
respect V. asius approaches Bridger material in the National Museum
referred to V. palustris, although M, is distinctly shorter. The depth
of the jaw, however, becomes a little shallower beneath the anterior
premolars and, as shown in No. 19336 as well as the type, the sym-
physeal portion is not so deep as in V. canavus.

CONDYLARTHRA
HYOPSODONTIDAE ”°
HYOPSODUS WORTMANI Osborn

There are in the upper Knight collection over 100 specimens be-
longing to species of Hyopsodus. Of these, 29 can be clearly determined
as belonging to Hyopsodus wortmant. Osborn (1902, p. 185) gave
the length of the lower molar series as varying from I1 to 13 mm.,
whereas Matthew (1915b, p. 317) in his key to Hyopsodus cited
10 mm. Probably neither are correct as Osborn likely included some
material which Matthew would have placed in Hyopsodus mentalis
lysitensis and Matthew may have derived the figure of 10 mm. as
characterizing the lower molar series from Osborn’s dimension for
the length of the upper molar series. In the few upper Knight speci-
mens in which all the lower molars are preserved the range is from
10.5 to 11.5 mm. An upper molar series measures 10 mm.

The teeth in the upper Knight materials regarded as representing
H. wortmani are markedly progressive; exhibiting well-developed
hypocones in the upper molars, the trigonid of the lower molars has
lost the paraconid, and the metaconid and talonid of P, are in an
advanced stage of development. The advance is noticeable over Gray

20 The use of the family name Hyopsodontidae appears to have become estab-
lished through usage, but largely as a result of a series of errors. Hyopsodidae
was first used by Schlosser in 1887, but in 1889 Lydekker used Hyopsodontidae,
attributing it to Schlosser with the comment “wrongly given as Hyopsodidae.”
Osborn in 1902 used Hyopsodontidae, also attributing it to Schlosser, and was
followed by Weber, and others; however, Hay (Bibliography) in 1902 and
Palmer (Index Generum Mammalium) in 1904 correctly credited Hyopsodidae
to Schlosser (1887) and Hyopsodontidae to Lydekker (1889), and the earlier
spelling was used by Wortman in his revision of the primates in 1903 and by
Loomis, with the correct source cited in his revision of Hyopsodus in 1905.
This was ignored by Matthew who reverted to Osborn in his revision of 1915.
In 1930 Hay used Hyopsodontidae but probably because he erroneously gave
the date of Lydekker’s spelling as 1887. Simpson, in his classification of 1945,
used Hyopsodontidae, but without reference to the earlier spelling.

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Bull materials, and in particular when comparison is made with the
more nearly equivalent-sized H. simplex.

In the New Fork materials is a fragmentary right mandibular
ramus with M;. The tooth corresponds well with equivalent material
of H. wortmani from the upper Knight beds.

HYOPSODUS MENTALIS (Cope)

The much larger series of specimens of Hyopsodus in the La Barge—
Big Piney collection are referred to the species Hyopsodus mentalis.
These include materials covering a rather noticeable range in size,
but within which no separation seems practical, or can logically be
maintained. The dimensions given by Cope (1875, p. 17) for Hyopso-
dus mentalis fall well within the upper limits of this material, but it
seems unlikely that the same species of this small mammal would be
represented in so widely separated localities. However, on the basis
of known material, the upper Knight form, represented by some 76
individuals, appears indistinguishable from the New Mexican species.
Moreover, the range of the length of the lower molar series is about 13
to 14.5 mm. This would appear to average less than the range 14 to 15
given by Matthew (1915b, p. 320) for the Lost Cabin and New
Mexican materials he included in the New Mexican species, and to
overlap the inferred range of his Hyopsodus mentalis lysitensis which
occurs in Lost Cabin as well as Lysite beds. Matthew’s figures here
should probably not be relied on as there are differences in cited
measurements between key and text for some of the lower Eocene
species of this genus.

Matthew (1915b, p. 319) has placed Hyopsodus lemoinianus Cope
in synonomy with H. mentalis, but it should be noted that the dimen-
sions for H, lemoinianus are at about the upper limit of the range, and
that the type of this species may have come from the older Gray Bull
beds. The species of Hyopsodus as currently recognized may not be
satisfactorily arranged; there is, however, abundant material repre-
senting this genus from various Eocene horizons, and this should
lend itself rather well to mathematical analysis.

HYOPSODUS BROWNI Loomis

Four specimens of Hyopsodus in the American Museum, collected
by Granger and Miller in 1906 from the Knight beds near Knight
station in southwestern Wyoming, do not appear to be conspecific
with any of the Hyopsodus material from the Big Piney—La Barge

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 61

region. Two of these, A.M. Nos. 12838 and 12839, are lower jaws
very close to material in the American Museum from the Lysite beds
identified as Hyopsodus browni. A third lower jaw, No. 12837, has
badly worn teeth which are somewhat smaller but may come within
the range of H. browni. The fourth, a maxilla with P* to M%, in-
clusive, is also somewhat smaller, corresponding to A.M. No. 12837.
The teeth in the maxilla, however, are larger than any in Hyopsodus
of the La Barge fauna, although an approach is made in one speci-
men, U.S.N.M. No. 19657, from southeast of Big Piney. Matthew
regarded Hyopsodus browni Loomis as a subspecies of H. powellianus,
between the latter and H. mentalis in size. He recorded (1915b,
p. 322) it as abundant in the Wind River Lysite, rare in the Big
Horn Lysite, and doubtful in the Lost Cabin. This form would appear
to be rather indicative of a Lysite age for the Knight beds on Bear
Creek.

PHENACODONTIDAE
PHENACODUS PRIMAEVUS Cope

The type specimen of this species was found by Cope in the type
Knight on Bear River but it appears not to be represented in upper
Knight beds of the upper Green River basin. There is, however, a
lower jaw portion in the National Museum collections which Dr.
D. H. Dunkle obtained in 1950 from lowest exposed Wasatchian beds,
just beneath variegated beds, in S$ sec. 11, T. 21 N., R. 117 W.,
about 3 miles east of Fossil in Lincoln County, Wyo. The specimen
includes P, and M,, and compares favorably with Phenacodus
primaevus.

MENISCOTHERIIDAE
MENISCOTHERIUM ROBUSTUM Thorpe
Plates 7, 8

Beyond doubt the most striking materials recovered from the upper
Knight deposits are the well-preserved skulls, jaws, and partial skele-
tons of the peculiar condylarth Meniscotherium, Remains represent-
ing Meniscotherium are nowhere so abundantly found, outside of the
locality of their original discovery in the lower Eocene of New
Mexico.

Following Cope’s discovery of a maxilla, to which he gave the name
Meniscotherium chamense (1874, p. 8), David Baldwin obtained con-
siderable material for Marsh and later for Cope from the New Mexi-

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

can or San Jose beds. The collection obtained for Cope was the basis
for an additional species, Meniscotherium terraerubrae Cope (1881b,
pp. 493-495), founded on a distinctly larger individual than M. cha-
mense, but since there seemed to be no clear separation of these two,
Granger (1915, p. 359) regarded M. terraerubrae as no more than
a subspecies of M/. chamense. Cope’s (1882b, p. 470) species, Menis-
cotherium tapiacitis, based on very much smaller material in the
Baldwin collection than either of the above, Granger regarded as
valid. Subsequent to Cope’s work on these, Marsh (1892, pp. 445-
449) named Hyracops socialis on material in Marsh’s Baldwin col-
lection, but a review of this species by both Granger and Thorpe
(1934, pp. 406-409) has resulted in its being placed in synonomy with
Meniscotherium chamense terraerubrae. A very considerable amount
of material representing Meniscotherium has subsequently been ob-
tained from the San Jose by Simpson and a thorough understanding
of these forms is anticipated.

Outside of New Mexico, materials from the Lost Cabin beds were
referred by Granger in 1915 to typical M. chamense and a form
recognized from Clark Fork Paleocene was described as Menisco-
thertum(?) priscum (1915, p. 360). Then in 1934 Thorpe described
a skull and jaw in the Marsh collection from 2 miles west of Aspen,
Wyo., which was collected by William Cleburne in 1875. To this he
gave the name Meniscotherium robustum. An examination of Veatch’s
(1907, pl. 3) geologic map, showing the position of Aspen as it was
originally located on the old route of the Union Pacific, shows M.
robustum type to have come from Knight beds to the east of the gen-
eral southward extension of the West Oyster Ridge structure and in
the Green River basin of Eocene deposition.

Material of Meniscotherium in the upper Knight beds of the upper
part of the Green River basin are equaled in abundance only by speci-
mens of Lambdotherium. Some 93 specimens, including eight skulls,
many jaws and maxillae, and various associated skeletal portions in this
collection appear to represent a single species which cannot be dis-
tinguished from the type of Meniscotherium robustum. In addition to
the type, and the Smithsonian collections from the vicinity of Big
Piney and La Barge, an M. robustum skull was found also by R. W.
Brown in a relatively thin zone of Knight beds exposed on Little
Bitter Creek between Rock Springs and Green River, Wyo. A de-
tailed description of the skull and other portions of the skeleton of this
Knight species will be omitted in deference to studies of related forms
in the San Jose fauna currently under way by Simpson.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 63

MENISCOTHERIUM, cf. CHAMENSE Cope

A surprising and rather significant discovery with respect to the
occurrence of Meniscotherium is that the material obtained from the
New Fork tongue represents a distinct species from the typical upper
Knight, M. robustum. Three individuals from separate localities along
Alkali Creek in Sublette County are all immature, but can be readily
compared with material of equivalent maturity belonging to the upper
Knight M. robustum. The New Fork specimens are seen to represent
a species distinctly smaller than M. robustum. The permanent anterior
upper molars preserved in two of the specimens are closely compar-
able in size, and apparently cannot be distinguished otherwise from
these teeth in the type of Meniscotherium chamense from the San
Jose lower Eocene beds of New Mexico.

PANTODONTA
CORY PHODONTIDAE
CORYPHODON, cf. RADIANS (Cope)
Plate 9

Fragmentary remains of Coryphodon are not among the more
common materials encountered during a search of upper Knight beds
in the vicinity of La Barge or Big Piney. Isolated teeth or tooth por-
tions representing some 13 individuals were preserved, although, to
my recollection, probably a much greater amount of very poorly
preserved or scarcely identifiable material was discarded or not col-
lected. However, a complete skull, U.S.N.M. No. 16701, in a rather
poor state of preservation, was found in SW4 sec. 24, T. 28 N.,
R. 114 W., ona branch of Dry Piney Creek in Sublette County, Wyo.
Attention was called to the occurrence of this specimen in the Knight
lower Eocene in a short paper concerned primarily with the first
collections obtained from the Almy Paleocene (Gazin, 1942, p. 217).
The skull was discovered in beds separated by faulting, as mapped by
A. R. Schultz (1914, pl. 1), from the general area of upper Knight
in the Green River basin. The locality was early regarded as possibly
somewhat older, and the preservation of the specimen together with
the lithology appeared nontypical; nevertheless, a jaw of Hyraco-
therium, cf. vasacciense was discovered on the same outcrop, support-
ing a lower Eocene age.

No less than 24 specific names and six generic names have been
applied to American Coryphodon, and revisions by Earle (1892),
Osborn (1898, pp. 188-218), and Matthew (1899, pp. 33, 36), while

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

differing in detail, have reduced the genera finally to one, and cut the
number of species almost in half. Four of the specific names pro-
posed, Coryphodon radians, Coryphodon semicinctus, Coryphodon
latipes, and Coryphodon hamatus, were based on materials from the
type Knight exposures, the oldest of which is Coryphodon radians
Cope (1872a), and which, incidentally, has priority over all others
in the North American Eocene.

M? in U.S.N.M. No. 16701 is slightly smaller than in the type of
Coryphodon radians, as reported by Osborn (1808, p. 198), measuring
about 31 by 40 mm. instead of 32 by 43 mm. It differs in form from
the type in the absence of a posterior spur from the metacone, the
presence of which was believed to distinguish C. radians. The length
of the premolar-molar series, about 160 mm., is intermediate between
that of Coryphodon hamatus and Coryphodon elephantopus. It differs
most noticeably from C. hamatus Marsh (1884, p. 52, fig. 55) in the
absence of the distinctive hypocone development seen in the upper
molars of Marsh’s type. C. elephantopus, as represented by U.S.N.M.
No. 111, cited by Osborn as a cotype, has transversely narrower upper
premolars than the Dry Piney Creek specimen. Without attempting
to revise or determine synonymy of Coryphodon species, a task of
major proportions, the specimen is referred tentatively to C. radians.
Some separate teeth of another individual, found in Fogarty Draw,
near the Green River, NW3 sec. 20, T. 28 N., R. 111 W., include a
second upper molar which measures about 37 by 43 mm. and may
represent a larger species, but this is not certain.

DINOCERATA
UINTATHERIIDAE
Cf. BATHYOPSIS FISSIDENS Cope

A pair of lower jaws, U.S.N.M. No. 19990, including the greater
part of the left ramus with P; to M; and a smaller part of the right
ramus with portions of P, to Ms, of a uintathere was found in the
New Fork tongue of Knight on Alkali Creek about 10 miles above its
junction with the Green River. The jaws are rather poorly preserved
and the teeth, particularly in the right ramus, are noticeably checked.

Details of the teeth correspond very well to those of the type of
Bathyopsis fissidens Cope (188ia, p. 194) although those in the New
Fork specimen are better worn. The posterior crest of the talonid of
the molars is truly a crest and not cuspate as in Probathyopsis. The
teeth are close to the same size, but slightly larger than in Bathyopsis

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 65

fissidens. However, from the measurements given by Cope it would
appear that P, in the type is much smaller than in the New Fork
specimen. The length given is appreciably less than the width, a con-
dition not verified in the figures. The relative size of P, corresponds
more nearly to the illustration. It seems likely that Cope’s dimension
is in error and that the length of P, is 0.0150 M instead of o.o105 M.

In spite of the similarity of the teeth, marked divergence is noted
in the form of the lower jaw. Preservation of the bone is very poor
but there is no doubt that the New Fork specimen lacked the massive
inferior flange characterizing the type. Wear of the teeth is evidence
of greater maturity than the type, and there is some evidence that
the diastema back of the canine was longer ; nevertheless, the inferior
margin of the jaw would appear to be somewhat like that of the
smaller and more primitive Probathyopsis praecursor Simpson (1929,
p. 2). This striking difference of form would suggest generic, or at
least specific, difference; however, a dimorphic condition has been
postulated by J. A. Dorr (1952, p. 89) with respect to Probathyopsis
materials from Hoback Canyon, so that the possibility of such a situa-
tion in the upper Wasatchian materials should not be ignored. Pend-
ing possible future discoveries clarifying the relationships, the New
Fork specimen is tentatively referred to Bathyopsis fissidens and
regarded as possibly female.

PERISSODACTYLA
EQUIDAE
HYRACOTHERIUM INDEX (Cope)

The specimens of Hyracotherium in the La Barge fauna readily
fall into three fairly well-defined groups, those of the smallest species
belong almost certainly to Hyracotherium index. Approximately 35
specimens are in this group including jaws and maxillae which give a
good representation of the cheek-tooth series.

Measurements of M, in seven lower jaws having this tooth show a
range of 6.3 to 6.8 mm. in length and 4.5 to 4.8 mm. in width with an
average of 6.6 and 4.6 mm., respectively. M, in the type of Hyraco-
therium index measures 6.7 and 4.7 mm.; a close correspondence. The
probability of identity would appear to be increased by the fact that
the type of H. index is also from the Knight formation, but at its type
locality on Bear Creek and hence evidently somewhat older. More-
over, it is interesting to note that H. index is the second species of
Hyracotherium to have been described from North America, the first
being Hyracotherium vasacciense and from the same beds.

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Two upper teeth representing Hyracotherium index are also in-
cluded in the Princeton University collection, No. 16172, from Knight
beds about 100 feet below Green River strata, 3 miles northwest of
Fossil, Wyo. These compare very favorably with H. index material
from the vicinity of Big Piney and La Barge.

HYRACOTHERIUM VASACCIENSE (Cope)

A much larger number of specimens, approximately 67, from the
upper Knight are in a size group clearly distinct from that of H. index.
The range in size of M, in 14 jaws, in which this tooth is present and
certainly determined, is 7.4 to 8.4 mm. in length and 5.5 to 6.5 mm.
in width, with an average of 7.8 and 5.9 mm., respectively. The type
of Hyracotherium vasacciense consists of a jaw portion with only M,
preserved. This tooth measures 8.2 mm. in length and 6.0 mm. in
width across both trigonid and talonid, a striking correspondence and
well within the range. Granger (1908, p. 222) was inclined to sup-
press this name but I find it advisable to revive it as the most logical
and applicable designation for the rather large series of specimens
from these beds. Suppression of this name was awkward inasmuch
as there is no question of its representing Hyracotherium and it is the
oldest for North American species. So far as being indeterminate,
the Knight collections under consideration would appear to give it
definition and practical significance.

In addition to the above collection from upper Knight beds of the
upper Green River basin, there are three right upper molars (A.M.
No. 12841) of one individual from the Knight on Bear River near
Knight station in the American Museum collection which are about
the right size and surely represent Hyracotherium vasacciense.

HYRACOTHERIUM, cf. VENTICOLUM Cope

Four specimens in the upper Knight collection appear rather large
to be included in the group identified as Hyracotherium vasacciense.
They do not include M;, but other teeth are found to be 10 to 20
percent larger on various selected measurements than some of the
large individuals of H. vasacciense. These unusually large specimens
are tentatively referred to the Lost Cabin species, Hyracotherium
venticolum Cope (1881a, pp. 198-199).

A left maxilla with P., Py, and M, in the Princeton collection,
No. 16173, from near Fossil, Wyo., first thought to be tapiroid, is
probably also to be referred to Hyracotherium venticolum. The P.
shows a prominent anterolingual crest, but with little evidence of

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 67
accessory cuspules. It resembles P* in Homogalax, but can be
matched in fourth premolar material in the upper Knight collection
from the Green River basin referred to Hyracotherium. The first
molar is more equine than tapiroid and is certainly not Heptodon.

Isolated upper and lower teeth in the New Fork collection compare
favorably with Hyracotherium venticolum. Various measurements
of these teeth are very close to those in the type of this Lost Cabin
species.

HYRACOTHERIUM, possibly H. CRISTATUM Wortman

Some lower teeth (A.M. No. 12842) among the materials collected
by Granger from the type Knight appear too large to be included in
H. vasacciense or H. venticolum and may represent one of the larger
species such as H. cristatum. They are too narrow to belong to the
Wind River H. craspedotum.

BRONTOTHERIIDAE
LAMBDOTHERIUM POPOAGICUM Cope
Plate 10

As many specimens of this earliest known titanothere were found in
the upper Knight beds as of Meniscotherium, and several were ob-
tained from the New Fork; however, the material for the most part
was more fragmentary than that of Meniscotherium. Nevertheless,
one of those rarities in collections of Lambdotherium, an almost com-
plete skull, is included.

There is surprisingly little published information on the skull of
Lambdotherium. The reconstruction in Osborn’s (1929, fig. 233)
monograph on the titanotheres is based on a maxilla and jugal, a
portion of the top of the cranium, the zygomatic process of the
squamosal, and a condyle. U.S.N.M. No. 19761, from a locality 12
miles north of Big Piney, Wyo., though somewhat crushed and dis-
torted, in essential details lacks only the basicranium. The rostrum
of this skull is broad and the nasals strong and protruding. As noted
in Eotitanops borealis (Cope) (see Osborn, 1929, fig. 251) the skull
is very broad between the orbits, and the postorbital processes are
prominent and partially enclose the orbit posteriorly. The most notable
difference from Eotitanops borealis, in dorsal aspect, is that the
sagittal crest extends much farther forward, and the anterior margins
of the temporal fossae curve more forward as well as inward from the
postorbital processes to meet in the sagittal crest. In Eotitanops and

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

later genera the anterior portions of the temporal fossae are widely
separated and are divided by a sagittal crest only in their posterior
parts. By virtue of this difference the postorbital processes of the
frontals are directed backward in Lambdotherium instead of forward
as in Eotitanops and others.

The equine occiput in posterior view is high, narrow, and some-
what rectangular above the mastoid portion, and is directed backward
rather than being erect as in Osborn’s figure of Gregory’s restoration.
Two rather prominent vascular foramina are noted in the antero-
ventral portion of the occiput; apparently the forward of these is
between the periotic and squamosal and the posterior and larger
between the periotic and exoccipital, somewhat as in horses.

In the absence of the basicranial portion, little of importance, other
than the dentition, is observed in the ventral view of the skull. It is
noted, however, that the posterior narial opening extends forward
between the tooth rows to a point about even with the posterior
margin of M?. Also, the postglenoid process of the squamosal is not
nearly so prominent and the circular canal for the audital tube is not
so nearly enclosed below by the squamosal from behind.

A study of the dentition together with a revision of the known
species of Lambdotherium was made by Bonillas (1936) on the basis
of materials in the California Institute of Technology from Knight
beds between La Barge and Big Piney. Little need be added to his
description of the permanent teeth, except perhaps to comment on
the character of the so-called mesostyle and other details of P* and
P*, The nearly median external style is not in the true position of a
mesostyle but appears to be a continuation onto the cingulum of the
external rib of the metacone, posterior to the position of the meso-
style between the paracone and metacone seen in the molars. It is
weak or absent on these teeth. Also the transverse lophs of P* are
variable and may or may not exhibit intermediate cuspules. More-
over, Bonillas noted a strong internal cingulum on P*, but this is
variable, as in the molars, and may be discontinuous,

Several milk dentitions are included in the Knight collection which
call for some comment. Dp’ appears to be identical to this tooth in the
permanent series and may indeed be the same, although it is in a
position immediately anterior to Dp* without the diastema of the
permanent series. Dp* to Dp* are, of course, decidedly more pro-
gressive than their counterparts in the replacing series. Dp? is very
much like P? except for sharper, better-developed cusps. Dp* and
particularly Dp* are molariform. In Dp* the cusp having the position

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 69

of a protocone, as in other perissodactyles of the Eocene, is not so
lingual in position as the hypocone, and the transverse lophs with their
accessory cuspules, are not developed. However, the crown of Dp*
appears to be identical in every respect, except smaller size, to M?.
Both of the posterior upper milk teeth have fully developed meso-
styles.

In the lower series Dp, is an elongate tooth with a distinct para-
conid and metaconid, but scarcely more than a prominent cingulum
and a buttresslike rugosity posteroexternal to the metaconid, repre-
senting the talonid. The trigonid of Dp; is also elongate and the three
cusps are almost equally developed ; however, no metastylid was noted.
The talonid of this tooth is molariform; broad and basined with a
well-developed hypoconid and entoconid. Dp, is smaller than M,,
but apparently molariform in all respects.

HELALETIDAE
HEPTODON, cf. VENTORUM (Cope)

Remains of Heptodon from the upper Knight in the Big Piney—
La Barge area are next in abundance to those of horses; however,
among the 30-odd specimens none are outstanding or particularly
comment-worthy. They consist for the most part of jaw fragments
with one to three teeth preserved. One jaw (U.S.N.M. No. 19784),
however, is almost entire and although only P, and P; are preserved
the alveolar or root portions of all the remaining teeth from the in-
cisors to M; can be seen and over-all proportions determined.

No. 19784 shows a single small alveolus for P, immediately anterior
to P,. The presence or absence of P, is probably not significant
although Seton (1931, p. 47) noted its presence in Heptodon browno-
yum in comparison with other species. The diastema between the
canine alveolus and that for P, is about 22 mm. The length of P,
to P,, inclusive, is about 22.5 mm. and for M, to M; about 38.5 mm.
These proportions are distinctly less than in Heptodon posticus or
Heptodon brownorum and somewhat greater than in Heptodon calci-
culus. The correspondence is closest to Heptodon ventorum although
measurements of teeth in the type are greater than in most of the
Knight specimens. There is some variation in size of teeth between
specimens in the collection, but I am inclined to regard them as rep-
resenting a single species which may well be H. ventorum.

Among the specimens collected by Granger and Miller from beds
in the vicinity of the type locality of the Knight formation is a frag-
ment of a tapiroid lower jaw (A.M. No. 12840) with P3, P,, and the

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

anterior portion of M,. Like the upper Knight materials from the
vicinity of Big Piney and La Barge, the teeth are a little smaller than
in the type of Heptodon ventorum but apparently not closer to H.
calciculus. The better development of the entoconid on P,, moreover,
suggests H. ventorum rather than H. calciculus as represented by
their types.

A single lower tooth in the Princeton material, No. 16174, from
near Fossil, Wyo., can be almost exactly duplicated by Dp, in Hepto-
don material from the upper Green River basin and in American
Museum Heptodon material from the Wind River basin.

HEPTODON, species

In the New Fork collections are various isolated teeth and tooth
portions, including an M, which is a little larger than in the upper
Knight material referred to Heptodon ventorum. The M3; has about
the length and width of this tooth in the type of Heptodon posticus;
however, on this basis it cannot be clearly distinguished from H.
brownorum.

HYRACHYIDAE
HYRACHYUS, species

A complete last upper molar and fragments of other teeth repre-
sent a species of Hyrachyus in the New Fork fauna not far in size
from Hyrachyus agrestis, or H. modestus as revised by Wood, from
the Bridger middle Eocene. McGrew (in Donovan, 1950, p. 64)
recognized the presence of Hyrachyus in the New Fork on the basis
of tooth fragments collected by J. H. Donovan, a student at the Uni-
versity of Utah, however, in the absence of other, more critical evi-
dence, regarded the beds as of Bridger age. Hyrachyus has been
found, though rarely, in Lost Cabin beds. The New Fork M* meas-
ures 20.3 mm. anteroposteriorly by 22.9 mm. in its greatest transverse
diameter.

ARTIODACT Yi”:
DICHOBUNIDAE
DIACODEXIS, near D. SECANS (Cope)
A single specimen, U.S.N.M. No. 19218, from north of Big Piney,
consisting of lower jaw portions and including M, from both right
and left sides, and left M3, is regarded as belonging to a species of

Diacode.xis close in size to Diacodexis olsen Sinclair (1914, pp. 292-
293) or Diacodexis secans (Cope) (1881a, pp. 187-188). M, is the

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN Fifa

same length as the tooth in D. secans or D. olseni but is appreciably
narrower than in the latter. Mz; in Sinclair’s type is not complete so
that the length cannot be determined, but its width is also much greater
than that in No. 19218. M; in D. secans is only a little wider and
longer. The length and greatest width of M, in the Knight specimen
are 4.6 and 3.4 mm., respectively. The dimensions for Mz; are 5.5
and 3.7 mm. There is a likelihood that the Knight specimen repre-
sents Diacodexis secans although it is noted that the apices of the
cusps in the type are noticeably closer together transversely, and the
entoconid of M; is much weaker. Moreover, in the absence of P,,
which is surely distinctive of D. secans, no certain assignment can be
made.

Sinclair’s revision of Diacodexis brought some order out of chaos,
but left something to be desired in omitting consideration of Diaco-
lexis nuptus and Diacodexis secans. Also, from a consideration of
the measurements of the type of Diacodexis chacensis and other
material from the San Jose beds, together with a large suite of speci-
mens from the Gray Bull, I am inclined to regard the Gray Bull
species as specifically distinct from Diacodexis chacensis. The speci-
men that Sinclair referred to as a paratype of D. chacensis did not
come from the San Jose and is appreciably smaller than the type, as
measured by Cope (1875, p. 15). The position of the type specimen
of D. chacensis at the upper limits, or distinctly beyond the upper
limits of Gray Bull materials, in various measurements, coupled with
geographic and horizon differences, would lead one to regard the
species as distinct. For this reason I would prefer to revive the oldest
Gray Bull name, Diacodexis metsiacus (Cope) for these specimens.
Diacodexis brachytomus (Cope) and Diacodexis laticuneus (Cope)
are probable synonyms. Diacodexis robustus Sinclair from the Gray
Bull is apparently valid. Diacodexis nuptus (Cope) and Diacodexis
secans (Cope) are described from the Wind River beds, hence, ac-
cording to Simpson (1933, pp. 115-116), are from the Lost Cabin
horizon. D. nuptus is exceedingly small in comparison with Gray
Bull materials, and the paraconid in the preserved lower molar is
rather well forward in position and slightly less lingual than observed
in the Gray Bull species. D. secans, on the other hand, has large
teeth, and is particularly characterized by the relatively large size of
P,. It is further observed that the molars in D. secans have the apices
of their cusps relatively closer together transversely than in any other
Diacodexis material observed. Diacodexis olseni, as noted above, is
characterized by the relatively greater width of the molars, No data

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

exist on the variation in this respect, nor in the characters of D.
secans. It is noted that the difference in length of P, between D. secans
and D. olseni is a little greater than Sinclair has shown as the varia-
tion of the length of this tooth in Gray Bull materials.

DIACODEXIS?, species

Van Houten (1945, p. 458) noted the presence of Diacodexis
material from the Knight formation in the collections of the American
Museum. Two lower jaws are so recorded in their catalog but these
could not be found. They were collected by Granger and Miller from
vicinity of the type Knight section on Bear Creek, near Knight sta-
tion. There is no evidence as to the species represented.

BUNOPHORUS, cf. MACROPTERNUS (Cope)

Difficulty was early encountered in attempting to identify, as rep-
resenting Bunophorus or Wasatchia, certain materials in the upper
Knight or La Barge collection. Review of the dental characters, out-
lined by Sinclair (1914, pp. 268-276) as distinguishing these two
genera, would appear to be resolved into inflation of the lower pre-
molars, the extent to which the paraconid is developed on lower
molars, and the form of M;. The premolar difference is largely one
of degree, the presence or absence of a weak or vestigial paraconid
is not always significant in other groups, and the form of Ms; is likely
variable as in Diacodexis. However, the combination of the characters
outlined by Sinclair, together with the development of the hypo-
conulid, which was noted to be more distinctive in those specimens
exhibiting a more distinctive paraconid, might justify retention of
both genera.

The upper Knight collections include five lower-jaw portions very
close to one another in size and form of teeth, and surely represent
a single species. The best of these is a right mandibular ramus,
U.S.N.M. No. 19210, with P, to Ms, inclusive. The teeth in this are
moderately worn so that the presence or absence of a paraconid cannot
be determined; however, P, would appear to be somewhat more
inflated than in figures of Wasatchia. M, and M3; in No. 19208 are
slightly worn and do not appear to have had a paraconid, and M;
conforms to the description of Bunophorus except that the talonid
is nearly as wide as trigonid. In No. 19209 P, is a little larger and
more inflated than in No. 19210, and M, shows a very small enamel
lake representing the paraconid. M, in No. 19207 also shows definite

No. 18 LOWER EOCENE KNIGHT FORMATION

GAZIN 73

evidence of a paraconid, whereas M, does not. However, Mz in No.
19213 shows a small paraconid but is otherwise not distinctive. Sin-
clair observed the presence of a small paraconid on M, of Bunophorus
macropternus and a vestige on M; but not on My. The upper Knight
material, though limited, lacks a vestige of this cusp on Ms, but in
one instance includes it on M,. The correspondence of the upper
Knight form would appear to be closer to Bunophorus than to Wasat-
chia and in size is very near Bunophorus macropternus.

In addition to the lower jaws there are two specimens showing
upper teeth, believed to be of the same species. One of these, U.S.
N.M. No. 19212, includes M* and M? and the other is an isolated
molar, probably M*. Upper teeth of Bunophorus have not previously
been described, and as may be anticipated, their resemblance is very
close to those figured of Wasatchia. The upper Knight superior teeth
are relatively little worn and the most striking feature is their ex-
tremely bunodont appearance. The cusps are all blunt, nearly circular
cones, with but slight buttresses on the anterior and posterior slopes
of paracone and metacone, and between the protocone and the two
accessory cusps, the protoconule and metaconule. There is no great
disparity in size of cusps, although the protocone is slightly larger
than the about-equal paracone and metacone. Slightly smaller than
the latter two are the approximately equivalent protoconule, meta-
conule, and hypocone. The hypocone is at the posterolingual ex-
tremity of the cingulum, which surrounds all but the lingual surface
of the tooth.

HEXACODUS,?! new genus

Generic characters——Tooth structure much as in Sarcolemur Cope,
but P, appears more inflated medially with metaconid incipient or
weak. Trigonid of lower molars with inner pair of cusps higher than
protoconid and anteroposteriorly compressed, but less so than in
Sarcolemur. Paraconid and metaconid less closely appressed than
Sarcolemur, and metaconid rather than paraconid the larger and
higher. Cross crests of trigonid weak and oblique crest from hypo-
conid less distinctly developed on posteroexternal slope of metaconid.
Hypoconid crescentic and entoconid conical with hypoconulid on
posterior cingular shelf, distinct and decidedly posterior to hypoconid
and entoconid, as in Sarcolemur.

Type species —Hexacodus pelodes, new species.

21 From Greek éé, six; 4x7, point ; 650vs, tooth—with reference to the six cusps,
including the hypoconulid, of the lower molars.

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

HEXACODUS PELODES,??2 new species
Plate 11, figures I-3

Type.—Left ramus of mandible with P, to M2, inclusive, U.S.N.M.
No, 19215.

Horizon and locality—Upper Knight beds, La Barge fauna, SW4
sec. 33, T. 32 N., R. 111 W., 12 miles north of Big Piney, Sublette
County, Wyo.

Specific characters—tTeeth slightly smaller and relatively a little
narrower than in Sarcolemur pygmaeus. Other characters not dis-
tinguished from those of genus.

Description.—Eleven specimens, all lower-jaw portions or lower
teeth, belong to Hexacodus pelodes. The material was first thought
to represent Diacodexis, but upon more critical examination it was
found that only one specimen in the upper Knight collections of the
upper Green River basin represents Diacodexis. The remainder of
these smaller dichobunids are recognized to be of a form more closely
related to Bridger Sarcolemur.

The fourth lower premolar in the type of H. pelodes is perhaps a
little more rounded or inflated externally than in S. pygmaeus but the
metaconid, well developed in the latter, is incipient or scarcely more
than a flexure or swelling on the lingual side of this tooth in the
earlier type. The paraconid as in Sarcolemur is prominent and turned
sharply inward. On the rather small heel of P, in the H. pelodes
type a distinctly isolated entoconid is observed. Two other specimens
of H. pelodes exhibit Py, and in one this tooth is more inflated and
the accessory cusps are more obscure. In the other, the tooth is only
slightly wider and has the equivalent parastyle, a better-developed
metaconid, but apparently no entoconid.

The lower molars of H. pelodes are less specialized in the peculiari-
ties of the trigonid noted in Sarcolemur pygmaeus, although the talo-
nids are very similar in the two forms. In S. pygmaeus it is noted that
the arcuate anterior crest of the trigonid may be somewhat higher
than the posterior, and the paraconid is set slightly inward and a little
higher than metaconid. Both are higher than the protoconid and these
differences in height are further emphasized by the manner in which
the tooth is worn through occlusion. Moreover, in Sarcolemur the
metaconid appears to be joined from near its summit to the anterior
wing of the crescentic hypoconid. In some third molars the metaconid-

22 From Greek m7\wdns, muddy; with reference to Muddy Creek, where the
type specimen was found.

No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN 75

hypoconid crest is more conspicuous than that between the metaconid
and protoconid. In H. pelodes the paraconid and metaconid, though
high and close together, are not so closely appressed as in Sarcolemur
and the metaconid is a little the larger; however, in wear the meta-
cone may be reduced so that the tooth more closely resembles Sarco-
lemur. The crest from the hypoconid rises on the posterointernal
slope of the metaconid, but not so high and with less emphasis.

The talonid of the first two molars, as in Sarcolemur, exhibits a
crescentic, though high-cusped, hypoconid which likewise does not
connect directly with the prominent conical entoconid, but sends a
weak spur posteriorly from the horn of the crescent to the hypo-
conulid which is situated on a prominent posterior ledge or cingulum.
The structure of this portion of the tooth is entirely unlike that in
Diacodexis. In M; of H. pelodes the hypoconulid is an exaggeration
of the condition noted in preceding molars, and with this portion of
the heel transversely more constricted than in Diacodexis. A central
crest extends forward from the hypoconulid to the cleft between the
hypoconid and entoconid, much as in Sarcolemur.

Discussion—Hexacodus pelodes would appear to be the first au-
thentic record of the Sarcolemur or Antiacodon type of dichobunid
in the lower Eocene, although species from Wasatchian beds have
been referred to this group. Erroneously allocated have been Antia-
codon mentalis and Antiacodon crassus. The first of these is now
included in Hyopsodus, and the second does not represent this group,
as may be inferred from Cope’s illustration (1877, pl. 45, fig. 15)
and as he later recognized (1884, p. 233), although its correct alloca-
tion is not known. A third name, Sarcolemur bicuspis, was listed
without description by Loomis (1907b, p. 357) in a faunal list of
materials collected from the lower levels of Tatman Mountain in
the Big Horn basin. The name is a nomen nudem and there may be
some doubt as to whether the material which he so designated repre-
sented this genus. Simpson (1945, p. 143) has listed Sarcolemur as
lower Eocene and Antiacodon as middle Eocene, and although it
seems likely that the letter L, there used for lower Eocene, is a typo-
graphical error, a shadow of a doubt persists for the following reason:
The type of Sarcolemur is S. furcatus, the type specimen of which was
collected by Cope in 1872 “from the bluffs of the Upper Green River”
(1873b, p. 608). That year, according to Cope’s account (ibid.,
p- 545), he traveled up the Green River from about 17 miles above
the town of Green River, as far as La Barge Creek, so there is the
remote possibility that S. furcatus came from the upper Knight near

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I1I7

La Barge Creek. However, much of the material similarly labeled in
Cope’s collection is Bridger in age and none so far as known is Knight ;
moreover, the type of S. furcatus resembles other Bridger materials
rather than Hexacodus pelodes.

In all probability Sarcolemur is a synonym of Antiacodon. Cope
originally referred his species to Marsh’s genus, but later erected
Sarcolemur because he felt Marsh’s description was inadequate (1877,
p- 148). Antiacodon venustus Marsh (1872, p. 210) has never been
figured, nor the type, so far as I know, compared with that of Sarco-
lemur furcatus, so that the equivalence of these two genera remains
in doubt. Nevertheless, the pertinent Bridger materials in the Na-
tional Museum collections, apparently representing but a single species,
are cataloged as Antiacodon pygmaeus.

MEASUREMENTS OF TYPE LOWER JAW OF
Hexacodus pelodes, NEW SPECIES, U.S.N.M. NO. 19215

Menethrorib eto Wie, Ane. Gers. sicistets ecg are wie oe Suenera se ate crate neem 12.8
P., anteroposterior diameter: transverse diameter............ 4.5: 2.8
Ms, anteroposterior diameter: greatest transverse diameter.... 4.2: 3.0
M2 “ ‘ “ “ “ A.A: 39
Depth of jaw beneath point between M: and Mz, lingually.... 6.7

HEXACODUS UINTENSIS,?? new species
Plate 11, figure 4

Type—Left ramus of mandible with M, and M,, Princeton Uni-
versity collection No. 16175.

Horizon and locality—Knight beds, too feet below Green River
formation, about 3 miles northwest of Fossil, Uinta County, Wyo.

Description—Distinctly larger than Hexacodus pelodes with tri-
gonid relatively of somewhat greater anteroposterior extent. Para-
conid and metaconid more distinctly separated. Talonid much as in
A, pelodes with hypoconulid on posterior cingulum and distinctly pos-
terior to low divide between hypoconid and entoconid.

Discussion.—This species of Hexacodus from the Princeton locality
near Fossil is clearly distinct from that of the La Barge fauna, and is
somewhat more removed from the middle Eocene Sarcolemur in the
construction of its lower molars. In this respect it is a trifle more
generalized and less removed from the Diacodexis type of artiodactyl
molar.

23 Named for Uinta County, Wyo.

no. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN Fd

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No. 18 LOWER EOCENE KNIGHT FORMATION—GAZIN SI

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1937. Notes on the Clark Fork, upper Paleocene, fauna. Amer. Mus. Noyv.,
No. 954, pp. 1-24, figs. 1-6.
1945. The principles of classification and a classification of mammals. Bull.
Amer. Mus. Nat. Hist., vol. 85, pp. i-xvi, 1-350.
SINcLAIR, WILLIAM J.
1914. A revision of the bunodont Artiodactyla of the middle and lower
Eocene of North America. Bull. Amer. Mus. Nat. Hist., vol. 33,
art. 21, pp. 267-205, figs. 1-28.
SrncLarr, WILLIAM J., and GRANGER, WALTER.
1911. Eocene and Oligocene of the Wind River and Big Horn basins. Bull.
Amer. Mus. Nat. Hist., vol. 30, art. 7, pp. 83-117, figs. 1-4, pls. 4-9.
1912. Notes on Tertiary deposits of the Big Horn basin. Bull. Amer. Mus.
Nat. Hist., vol. 31, art. 5, pp. 57-67, figs. 1-2, pls. 5-6.
Tuorpe, MAtcotm R.
1934. Meniscotherium robustum sp. nov., and a discussion of Hyracops
socialis Marsh. Amer. Journ. Sci., vol. 27, pp. 401-409, figs. 1-6.
TROXELL, Epwarp L.
1923a. The Eocene rodents Sciuravus and Tillomys. Amer. Journ. Sci.,
vol. 5, pp. 383-396, figs. 1-23.
1923b. A new marsupial. Amer. Journ. Sci., vol. 5, pp. 507-510, figs. 1-4.
Van Houten, FRANKLYN B.
1945. Review of latest Paleocene and early Eocene mammalian faunas.
Journ. Paleont., vol. 19, No. 5, pp. 421-461, 1 fig., 4 tables.
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1907. Geography and geology of a portion of southwestern Wyoming.
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WILson, Rosert W.
1938. Review of some rodent genera from the Bridger Eocene. Amer.
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ington Publ. 584, pp. 67-164, figs. I-13.

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I17

Woop, Horace E., 2d, et al.
1941. Nomenclature and correlation of the North American Tertiary. Bull.
Geol. Soc. Amer., vol. 52, pp. 1-48, pl. I.
WorrMan, JAcoB L.
1903. Studies of Eocene Mammalia in the Marsh collection, Peabody Mu-
seum. Part II. Primates. Suborder Cheiromyoidea. Amer. Journ.
Sci., vol. 16, pp. 345-368, figs. 105-110, pls. 11-12.

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VOL. 117, NO. 18, PL. 1

SMITHSONIAN MISCELLAN EOUS COLLECTIONS

MARSUPIALS, INSECTIVORE. AND PRIMATES FROM THE
KNIGHT LOWER EOCENE

PLATE I

Fic. 1. Peratherium chesteri, new species: Right ramus of mandible (U.S.N.M.
No. 19199), type specimen, occlusal and lateral views. Approximately six
times natural size.

Fic. 2. Diacodon pineyensis, new species: Left ramus of mandible (U.S.N.M.
No. 19204), type specimen, occlusal and lateral views. Approximately four
times natural size.

Fics. 3 and 4. Peratheriwm edwardi, new species: 3, Left ramus of mandible
(U.S.N.M. No. 19200), type specimen, occlusal and lingual views; 4, left
maxilla (U.S.N.M. No. 19206), occlusal view. Approximately five times
natural size.

Fic. 5. Paratetonius? sublettensis, new species: Left ramus of mandible
(U.S.N.M. No. 19205), type specimen, occlusal and lateral views. Approxi-
mately five times natural size.

Fics. 6-8. Notharctus limosus, new species: 6, Left maxilla (U.S.N.M.
No. 19293), occlusal view; 7, right ramus of mandible (U.S.N.M.
No. 19294), type specimen, lateral view ; 8, left ramus of mandible (U.S.N.M.
No. 19294), type specimen, lingual view. Approximately three times
natural size.

PLATE 2

Fic. 1. Cynodontomys knightensis, new species: Left ramus of mandible
(U.S.N.M. No. 19314), type specimen, occlusal and lingual views. Approxi-
mately three times natural size.

Fries. 2-5. Undetermined stylinodont: Fore foot (U.S.N.M. No. 18425) ;
2, pisiform, lateral view; 3, scaphoid, proximal view; 4, 3d(?) digit, inner
view ; 5, fourth digit with unciform and fifth metacarpal, lateral view. Natural
size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 27, INOW 18; RE 2

INSECTIVORE AND TAENIONDONT FROM THE KNIGHT LOWER EOCENE

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO eii7/) NOS 6 Pies

TAENIODONT FOOT FROM THE KNIGHT LOWER EOCENE

Articulated right hind foot of undetermined stylinodont (U.S.N.M. No. 18425),
anteromedial view. Internal cuneiform and distal phalanx of 2d digit par-
tially restored from left foot. A portion of the proximal extremity of second
metatarsal not represented. Approximately four-fifths natural siz.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 18, PL. 4

TAENIODONT FOOT FROM THE KNIGHT LOWER EOCENE

Articulated right hind foot of undetermined stylinodont (U.S.N.M. No. 18425),
anterolateral view. Fifth metatarsal and distal phalanx of second digit partially
restored from left foot. Approximately four-fifths natural size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 18, PL. 5

CREODONTS FROM THE KNIGHT LOWER EOCENE

Fics. 1-3. Prolimnocyon clisabethae, new species: TI, Left maxilla (U.S.N.M.
No. 19348), occlusal view of M2 and M7’; 2, right maxilla (U.S.N.M.
No. 19348), occlusal view of P* and M’; 3, left ramus mandible (U.S.N.M.
No. 19350), type specimen, occlusal and lateral views. Approximately one
and one-half natural size.

Fic. 4. Vulpavus asius, new species: Left ramus of mandible (U.S.N.M.
No. 109337), type specimen, occlusal and lateral views. Approximately one
and one-half natural size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO ESI 7 NOMS REG

CREODONTS FROM THE KNIGHT LOWER EOCENE

Fic. 1. Uintacyon asodes, new species: Left ramus of mandible (U.S.N.M.
No. 19351), type specimen, occlusal and lateral views. Approximately one

and one-half natural size.
Fic. 2. Ambloctonus major Denison: Left ramus of mandible (P. U.

No. 14720), occlusal and lateral views. Natural size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS WoLs atil77, ING. webs be 7

MENISCOTHERIUM FROM THE KNIGHT LOWER EOCENE

Meniscotherium robustum Thorpe: Skull and mandible (U.S.N.M. No. 18283),
dorsal and lateral views. Upper, approximately three-fourths natural size;
lower, two-thirds.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 117, NO. 18, PL. 8

MENISCOTHERIUM FROM THE KNIGHT LOWER EOCEN=

Mentscotherium robustum Thorpe: Above, skull and mandible (U.S.N.M.
No. 19505), lateral view; below, skull (U.S.N.M. No. 18260), ventral
view. Approximately three-fourths natural size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 117, NO= ws REas

CORYPHODON FROM THE KNIGHT LOWER EOCENE

Coryphodon, cf. radians (Cope): Skull (U.S.N.M. No. 16701), ventral view.
Approximately one-third natural size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. L777 NOS Ls; RE Lo

GHT LOWER EOCENE

LAMBDOTHERIUM FROM THE KNI

Lambdotherium popoagicum Cope: Skull (U.S.N.M. No. 19761), dorsal, lateral
and ventral views. Approximately one-half natural size.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 217; NO: Ve; (PEs

ARTIODACTYLES FROM TH= KNISHT LOWER ECCENE

PLATE II

Fics. 1-3. Hexacodus pelodes, new genus and species: 1, Left ramus of mandi-
ble with Ps—M. (U.S.N.M. No. 19215), type specimen, occlusal and
lateral views; 2, right ramus of mandible with M; (U.S.N.M. No. 19217),
occlusal and lateral views; 3, right M; (U.S.N.M. No. 19219), occlusal

» and lateral views. Approximately four times natural size.

Fic. 4. Hexacodus uintensis, new species: Left ramus of mandible with
Mi-Mz (P.U. No. 16175), type specimen, occlusal and lateral views.
Approximately four times natural size.

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